Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Sex-ratio of Skylark Alauda arvensis broods in relation to timing of breeding

Capsule Earlier broods tend to be more male biased than later broods     

Reproductive success of polygynous male corn buntings (Miliaria calandra) as confirmed by DNA fingerprinting

Despite a sex ratio approximating to unity, female corn buntingswere not equally distributed among males. In 1989 and 1990,41.2% of 50 males were monogamously paired, 29.4% were polygynous,and 23.5% were unpaired. Polygynous males usually paired withtwo females, although in 1990 three males were trigamous. Polygynousmales fledged more offspring from their territories than didmonogamous males, mainly because they had more mates. The fledgingsuccess per nesting female was slightly higher in territoriesof polygynous males, but not significantly so. DNA fingerprintingwas used to confirm the true paternity of 44 offspring from15 broods and the true maternity of 50 offspring from 16 broods.A further 12 offspring from three broods for which neither putativeparent was available were also fingerprinted. Actual reproductivesuccess of parents was close to that inferred from observationsof number of young raised. There was only one brood, containingtwo chicks (4.5% of offspring, or in 6.7% of broods), wherethe chicks were not fathered by the male defending the territory.However, this nest was close to the territory boundary, andthe defending male may have been assigned incorrectly. Therewere no cases of intraspecific brood parasitism (n = 16 broods).The copulation rate was low, and extrapair copulation attemptswere rare, probably because of the poor chances of sneakingonto a neighbor's territory undetected and the costs of leavinga territory unguarded.     

Effect of cereal harvesting method on a recent population decline of Corn Buntings Emberiza calandra on the Western Isles of Scotland

Capsule Population decline since 1995 is associated with the harvesting of cereals as arable silage.

Aim To survey Corn Bunting populations on the Western Isles and test whether population changes since 1995 are associated with change in cereal harvesting methods.

Methods Twelve areas of machair on the Uists and Benbecula were re-surveyed in 2002, after previous surveys in 1983 and 1995. These and other areas of suitable habitat on these islands, plus Berneray, Baleshare, Vallay, Barra and Vatersay, were surveyed in 2003–05 to provide a breeding population estimate for the Western Isles. Cereal strips in the 12 core survey areas were mapped in 2002 and the timing and method of harvesting of each was recorded. Cattle-feeding stands in 24 crofting townships were visited in March 2003 to record the type of fodder available, measure grain abundance and ripeness in fodder samples, and record foraging Corn Buntings. Experimental trials tested whether Corn Buntings selected fodder with riper grain.

Results The Western Isles Corn Bunting population was estimated at 117 occupied territories in 2005, a decline of 17% since 2002/03, and of at least 62% since 1995. Population decline was greater in areas with higher proportions of the cereal area harvested as arable silage. Arable silage contained fewer and less-ripe grains than traditionally harvested cereal, and was less likely to attract feeding Corn Buntings. Corn Buntings preferred to feed on silage containing riper grain.

Conclusions The remaining Corn Bunting population of the Western Isles is declining rapidly, probably because of reduction in winter grain supply caused by change from traditional reaper-binder harvesting and stacking of fully ripe cereals to harvesting the crop as unripe or partially ripe arable silage. Management change to restore a supply of ripe grain over winter is likely to be essential if this population decline is to be halted and reversed.     

Agricultural habitat-type and the breeding performance of granivorous farmland birds in Britain

Conservation concern about granivorous birds has led to the implication of changing agricultural practices as causes of widespread population decline. We investigate relationships between breeding performance and the agricultural environment for ten granivorous farmland bird species (Stock Dove Columba oenas, Skylark Alauda arvensis, Tree Sparrow Passer montanus, Chaffinch Fringilla coelebs, Greenfinch Carduelis chloris, Linnet C. cannabina, Bullfinch Pyrrhula pyrrhula, Reed Bunting Emberiza schoeniclus, Yellowhammer E. citrinella and Corn Bunting Miliaria calandra). We analyse long-term, extensive data from the British Trust for Ornithology's Nest Record Scheme on breeding performance per breeding attempt with respect to farmland type (arable, grazing or mixed) and time (pre- and post-1975–76). The influence of habitat is investigated at two different scales: within the nesting territory and at the landscape level. Relationships between farmland type and (temporal changes in) breeding performance tended to be species-specific, but a few patterns were each common to some species. Improvements in breeding performance occurred across all three farmland types for four declining species. Grazing farmland seems to have deteriorated as breeding habitat for Linnet and arable/mixed farmland for Reed Bunting. Mixed farming at the territory scale supported better breeding performance for four species, three of which (Bullfinch, Yellowhammer and Corn Bunting) have declined concurrently with mixed farming. Pastoral landscapes supported better breeding performance for up to seven species, six of which have undergone large declines. Arable landscapes supported better breeding only for the stable or increasing Chaffinch and Greenfinch. Different relationships between farming regime and breeding performance were found at the two scales considered.     

Microhabitat use and social structure in Linnet Carduelis cannabina and Corn Bunting Miliaria calandra at a winter roost in central Spain

Capsule Both species selected particular patches for roosting, and Linnets but not Corn Buntings showed a social structure with males occupying the preferred roosting positions.

Aims To evaluate microhabitat preferences of Linnet and Corn Bunting within a winter communal roost, and to examine roost structure according to sex, age and body size.

Methods We captured Linnets and Corn Buntings along a habitat gradient in a reed-bed in central Spain when they were coming to roost. Each bird was sexed, aged, and several body traits were measured. We also characterized the habitat patches where the birds were trapped and grouped patches according to their similarity.

Results Three groups of patches were defined according to reed structure. Linnets preferred patches with low density of high and thick shoots, while Corn Buntings preferably used patches with intermediate values for these variables. Interspecific competition for preferred positions was not evident. Male Linnets were proportionately more abundant in the preferred patches, and individuals in lower body condition, independent of sex, age or body size, occupied the non-preferred patches. Male Corn Buntings were less abundant, bigger in size, and showed better condition than females, although differences in body size and condition were not apparent among patches within the roost.

Conclusion Our results suggest that among-patch differences in structure and location within a reed-bed generate microhabitats that differ in quality for roosting passerines. This habitat heterogeneity in turn seems to be responsible for the social spatial structure observed in Linnets, in which females and individuals in poor physiological condition were displaced, possibly through competition, towards non-preferred positions. The relatively small body size of Linnets compared to Corn Buntings might explain Linnets' preference for patches that seem to provide more protection against harsh weather conditions.     

Conservation insights from changing associations between habitat,territory distribution and mating system of Corn Buntings Emberiza calandra over a 20‐year population decline

Associations between habitat and animal distributions are widely used to make conservation recommendations. However, short‐term studies do not allow investigation of temporal variation in associations as habitats change or populations decline. Here we quantify changes in the habitat attributes and distribution of breeding territories in a multiple‐brooded, crop‐nesting farmland bird, the Corn Bunting Emberiza calandra, over a 20‐year period during which the study population declined by 91%. Corn Buntings were positively associated with weedy fields, overhead wires, spring cereals and, in early summer, with winter barley and forage grass. Territory associations with wires (positive) and fallow (positive in early summer, negative in late summer) were stronger in later years when the population was smaller. Trends in the proportions of males holding territories into late summer (decline), and mated polygynously or not at all (increases), suggested that habitat quality declined and became more spatially variable in later years. Field size increased and weed abundance within fields declined, reducing the availability of field‐boundary song‐posts, invertebrate‐rich foraging habitats and physical cover for nests within crops. Conservation recommendations are for weed‐rich or under‐sown spring cereals and winter barley combined with late‐cut hay and fallow, especially when offered close to song‐posts such as overhead wires. We also demonstrate the value of long‐term studies by comparing our 20‐year analysis with 3‐year periods at the start, middle and end of this period, and expose some risks of conservation recommendations derived from short studies.     

Problems of censusing Reed Buntings,Sedge Warblers and Reed Warblers

The daytime dispersal of Starlings from an urban winter communal roost was monitored by colour marking 1424 birds according to age and sex. Reports received from members of the public suggested that dispersal was homogeneous with respect to age and sex composition at all distances from the roost. There was evidence that dispersal was geographically heterogeneous with respect to sex composition, and this observation is discussed in relation to correlated variation in agricultural practices.     

Agricultural intensification and the collapse of Europe's farmland bird populations

The populations of farmland birds in Europe declined markedly during the last quarter of the 20th century, representing a severe threat to biodiversity. Here, we assess whether declines in the populations and ranges of farmland birds across Europe reflect differences in agricultural intensity, which arise largely through differences in political history. Population and range changes were modelled in terms of a number of indices of agricultural intensity. Population declines and range contractions were significantly greater in countries with more intensive agriculture, and significantly higher in the European Union (EU) than in former communist countries. Cereal yield alone explained over 30% of the variation in population trends. The results suggest that recent trends in agriculture have had deleterious and measurable effects on bird populations on a continental scale. We predict that the introduction of EU agricultural policies into former communist countries hoping to accede to the EU in the near future will result in significant declines in the important bird populations there.     

Eggs, egg formation and the timing of breeding

The eggs of birds contain all the nutrients necessary for the developing embryo. The female has to obtain these nutrients, and while in some species these may be gathered and stored over a period of time, in many species they seem to be obtained on a daily basis. Obtaining the calcium necessary for the eggshell may pose special problems for some species, which may have to give up valuable feeding time in order to search for calcium.
Egg size varies considerably, even within species, and the reasons for this are not clear. In the Great Tit Parus major , large eggs produce nestlings which have a higher chance of hatching, surviving to fledging and surviving after fledging. However, large eggs require the female to acquire more nutrients to form them, and large chicks may sometimes be at a disadvantage. The amount of energy required to form a larger egg often seems small unless the metabolic costs of egg formation are taken into account.     

The timing of birds' breeding seasons: a review of experiments that manipulated timing of breeding

Reproductive success usually declines in the course of the season, which may be a direct effect of breeding time, an effect of quality (individuals with high phenotypic or environmental quality breeding early), or a combination of the two. Being able to distinguish between these possibilities is crucial when trying to understand individual variation in annual routines, for instance when to breed, moult and migrate. We review experiments with free-living birds performed to distinguish between the 'timing' and 'quality' hypothesis. 'Clean' manipulation of breeding time seems impossible, and we therefore discuss strong and weak points of different manipulation techniques. We find that the qualitative results were independent of manipulation technique (inducing replacement clutches versus cross-fostering early and late clutches). Given that the two techniques differ strongly in demands made on the birds, this suggests that potential experimental biases are limited. Overall, the evidence indicated that date and quality are both important, depending on fitness component and species, although evidence for the date hypothesis was found more frequently. We expected both effects to be prevalent, since only if date per se is important, does an incentive exist for high-quality birds to breed early. We discuss mechanisms mediating the seasonal decline in reproductive success, and distinguish between effects of absolute date and relative date, for instance timing relative to seasonal environmental fluctuations or conspecifics. The latter is important at least in some cases, suggesting that the optimal breeding time may be frequency dependent, but this has been little studied. A recurring pattern among cross-fostering studies was that delay experiments provided evidence for the quality hypothesis, while advance experiments provided evidence for the date hypothesis. This indicates that late pairs are constrained from producing a clutch earlier in the season, presumably by the fitness costs this would entail. This provides us with a paradox: evidence for the date hypothesis leads us to conclude that quality is important for the ability to breed early.     

Eggs, egg formation and the timing of breeding

The eggs of birds contain all the nutrients necessary for the developing embryo. The female has to obtain these nutrients, and while in some species these may be gathered and stored over a period of time, in many species they seem to be obtained on a daily basis. Obtaining the calcium necessary for the eggshell may pose special problems for some species, which may have to give up valuable feeding time in order to search for calcium.
Egg size varies considerably, even within species, and the reasons for this are not clear. In the Great Tit Pants major, large eggs produce nestlings which have a higher chance of hatching, surviving to fledging and surviving after fledging. However, large eggs require the female to acquire more nutrients to form them, and large chicks may sometimes be at a disadvantage. The amount of energy required to form a larger egg often seems small unless the metabolic costs of egg formation are taken into account.     

The breeding ecology of Reed Buntings Emberiza schoeniclus in farmland and wetland habitats in lowland England

We studied the nesting and feeding ecology of Reed Buntings Emberiza schoeniclus breeding on farmland and wetland habitats along the Trent Valley in Nottinghamshire, England. Rank and emergent vegetation accounted for most nests and most foraging by provisioning adults. Caterpillars and spiders accounted for 70% of chick invertebrate prey and all broods were fed cereal grain or other vegetable matter. Variation in the abundance of key invertebrate prey across habitats accounted for the foraging preferences of adult buntings. Depredation was the main cause of nest failure, and survival of nests at the egg stage was positively related to the extent of nest concealment. A measure of total brood biomass was positively related to the abundance of key invertebrate prey within 100 m of nests. Rank and emergent vegetation provided Reed Buntings with greater nest concealment and a richer source of invertebrate prey than agricultural habitats such as set-aside, cereals and oilseed rape. The provision of rank and emergent vegetation on farmland is likely to increase the nesting opportunities and productivity of Reed Buntings in agricultural landscapes.     

Subject: Eggs, clutches and the timing of breeding

Research Notes on Avian Biology 1994: Selected Contributions from the 21st International Ornithological CongressGeneral Biology: Clutch Size and Time

Subject: Eggs, clutches and the timing of breeding     

Mating system and timing of breeding in Holarctic waders

In passerine birds polygamous species lay eggs later than monogamous species. Yom-Tov has hypothesized that this is because polygamous males do not provision their mates with food, thereby delaying egg-laying. An indirect prediction of this hypothesis is that there should be no difference in laving dates between monogamous and polygamous birds in which mate provisioning does not occur. This prediction was tested by examining egg-laying dates in sympatric wader species that have contrasting mating systems but in which food provisioning of mates does not occur. Polygamous waders laid eggs later than monogamous waders, however, both in a comparison involving only calidridine sandpipers and in a comparison involving sandpipers and plovers. Accounting for phylogeny did not affect the direction of the results, which therefore cast doubt on the validity of the 'mate provisioning' hypothesis and indicated that features common to both passerines and waders were more likely to explain the differences in egg-laying dates between mating systems. Mating system and parental care system were strongly related, with all polygamous species being uniparental and monogamous species being biparental. It is suggested that seasonally early food supplies are of relatively low abundance, constraining species to biparental care and monogamy. By contrast, seasonally late food supplies are more abundant, making uniparental care more successful, thereby creating greater opportunities for mate desertion and polygamy.     

The timing of breeding and wing-moult of four African Sturnidae

Wing-moult of the Cape Glossy Starling, Red-winged Starling, Pale-winged Starling and Pied Starling was examined primarily from specimens in southern African museums. Breeding data were obtained from nest record cards.
The Cape Glossy Starling breeds from October to March, with the moult period from December to May. There is no evidence of moult-breeding overlap in individual birds. The Red-winged Starling breeds from September to March, while the moult takes place between November and April, overlapping with the second broods. The Pale-winged Starling breeds from October to April and moults between November and May. The Pied Starling moults between November and April, while breeding varies regionally, occurring concurrently with moulting in some areas.