Ecogeographic correlates of morphometric variation in the Red-billed Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus

We estimated the magnitude of intraspecific geographic variation in the Red-billed Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus from the analysis of external measurements of museum specimens collected throughout the Palearctic ranges of the species. By means of univariate and multivariate techniques we tested the importance of climatic and geographic factors as potential agents that might have shaped Red-billed and Alpine Chough morphology, discussing the dynamics of the interplay between natural selection, gene flow and phylogenetic constraints. Both species exhibited concordance in character differentiation. Bergmann's rule offered a valid explanation of Red-billed and Alpine Chough body size variation with the largest birds being found at higher elevation, or in colder and more arid regions. Shape was also important. The extremities of the body (bill and tarsus) were longer in warmer areas, in line with Allen's rule. In the Alpine Chough, temperature seemed to be the most important cause of body trait variation, whilst in the Red-billed Chough geographic distance among populations (a measure of geographic isolation) and altitude were also significant. In this paper we critically evaluate the validity of subspecific categories quoted in the literature, given that most variation appeared to be clinal.     

Linking territory quality and reproductive success in the Red-billed Chough Pyrrhocorax pyrrochorax: implications for conservation management of an endangered population

As changes in land use have been identified as the main factor explaining the decline in Red-billed Chough Pyrrhocorax pyrrochorax populations across western Europe, a study was carried out in Ouessant (western France) in order to assess the relationship between territory quality and reproductive success. As such an approach could be hindered by the fact that the birds' reproductive performance could be influenced by their breeding experience, we analysed both inter- and intranest-site variation in fledging success. Territory quality was quantified, combining habitat selection, territory size, the amount of feeding area and distance between nest and feeding area. Feeding habitats selected positively by Red-billed Choughs were characterized by a mean sward height of less than 5 cm. Foraging area was on average 21 557 m². Feeding areas (i.e. feeding habitat within foraging area) close to the nest were used preferentially. Fledging success appeared to be influenced neither by the total area of feeding sites in a Chough territory, nor by the mean feeding flight distance. However, fledging success adjusted to nest-site and year appeared to be influenced by feeding area close to the nest: one additional fledgling was expected for each additional 10 000 m² of feeding habitat within 300 m of the nest. These first results allowed us to consider recommending landscape management measures to ensure a favourable conservation status of local Chough populations.     

A comparative analysis of the foraging behaviour of the Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus coexisting in the Alps

The foraging behaviour of the Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus was studied during summer and autumn in the western Italian Alps. We assessed feeding times, feeding rates and foraging techniques associated with different foraging habitats. The Alpine Chough is mainly a ground surface feeder, stays for a rela-tively short time at a feeding site (on average 2.1 min) and feeds quickly (on average 9.2 items/min). In contrast, the Chough is almost exclusively an undersurface feeder (digger and prober), stays at a feeding site twice as long as the Alpine Chough (5.4 min) and feeds four times as slowly (2.2 items/min). These differences suggest that the degree of actual ecological overlap is almost as low as possible for two species using the same Alpine pastures as foraging sites. Interspecific coexistence has occurred through a clear differentiation of foraging strategies and diets. The foraging efficiency (in terms of feeding rate)of the Alpine Chough was always higher than that of the Chough in all habitats where they occurred together. The Alpine Chough was more variable in the use of foraging techniques and more diversified in the use of foraging habitats than the Chough. In both species, juveniles fed less efficiently than adults; the foraging behaviour of the Chough is probably more difficult to learn than that of the Alpine Chough. Other data have also shown that the Alpine Chough is more opportunistic than the Chough in using seasonally available food. Considering the above, we suggest that the foraging behaviour of the Alpine Chough is more flexible and, perhaps, better adapted to the high mountain Alpine environment than that of the Chough. Two hypotheses concerning the ultimate reason why the Alpine Chough and the Chough have evolved divergent beak morphology are discussed.     

A comparative analysis of the diets of the Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus coexisting in the Alps

The diets of the Chough Pyrrhocorax pyrrhocorax and the Alpine Chough Pyrrhocorax graculus coexisting in the western Italian Alps have been compared by faecal analyses. A total of 1581 fresh droppings (405 of the Chough and 1176 of the Alpine Chough) were collected in the pastures of the Rhêmes Valley, Aosta, Italy, from June to November 1992 and analysed in the laboratory, and a mean volume percentage for each item was calculated. Both species were omnivorous, including animal, vegetable and mineral dietary fractions. There was no overlap in the vegetable part of the diet since the Chough fed virtually exclusively on Yellow Gagea Gagea fistulosa bulbs (dug out from the soil), which were not taken at all by the Alpine Chough. Conversely, the Alpine Chough fed on berries and hips from September to November, but these were virtually ignored by the Chough. Even though animal items were collected by both species, their preferences were different. In June, Alpine Choughs largely collected cranefly (Tipula) larvae whilst Choughs mainly fed on Lepidoptera larvae. From July onward, Alpine Choughs mainly consumed grasshoppers while Choughs also collected Tipula pupae, Lepidoptera and fly (Bibionidae) larvae and beetles (Scarabaeidae and Staphylinidae). Interspecific morphological and behavioural differences may be partly responsible for the segregation observed. Chemical composition and caloric contents of the food items suggest that the balance between costs of collecting and benefits of consuming may also contribute to diet differentiation. Both species took a broad spectrum of food, and there was no clear indication that the different population densities of the two Choughs in the Alps were directly correlated with diet, even though some data suggest that during autumn the Alpine Chough might have a diet better adapted to the high mountain environment than the Chough.     

Effects of nest-site availability and distribution on density-dependent clutch size and laying date in the Chough Pyrrhocorax pyrrhocorax

We compared the breeding phenology and clutch size of Choughs Pyrrhocorax pyrrhocorax in three Spanish areas that differed in feeding habitat, breeding environmental conditions and the distribution and availability of different nesting sites (cliffs or artefacts). The variation in the timing of laying seemed to be related to differences in climatic conditions determined by the altitude. The variation in clutch size was associated with the different use that Choughs made of the feeding areas because of the contrasting availability and distribution of their nesting sites. Within southeastern Madrid, clutch size of the Chough decreased and its variance increased at high nest densities on cliff sites, suggesting that density dependence in fecundity arises from habitat heterogeneity rather than by interference. At high densities, individuals, territories, nest sites or a combination of these may differ in quality, thus promoting differences and increasing the variance in the initial investment of reproductive effort. The syncronization of the onset of laying at increased breeding densities suggests that social influences arising from communal foraging or avoidance of predation by early warning and predator swamping may be acting also. We suggest that nest-site availability and distribution have a major influence on the social organization of Choughs through their breeding and foraging strategies.     

Vocalizations and morphology: interpreting the divergence among populations of Chough Pyrrhocorax pyrrhocorax and Alpine Chough P. graculus

Capsule Differences in vocalizations among populations are mostly explained on morphological bases, but historical factors may have played a significant role in differentiation processes.

Aims To investigate the relationships among vocal and morphological variation in two corvids: Chough and Alpine Chough.

Methods We used data from 11 populations of Chough and seven populations of Alpine Chough spanning the Palearctic distribution of the two species. Three data sets (morphometry, spectrotemporal parameters of trill calls and acoustic repertoire) were analysed and their variation compared with uni- and multivariate techniques.

Results In both species, morphological differences among populations were correlated to spectro-temporal variation of trills; in particular, frequencies of calls were negatively correlated to wing length (an indicator of body size). By considering only co-existing populations of the two species, the magnitude of morphological and spectrotemporal divergence was similar.

Conclusions In both species, birds from populations with similar morphology uttered similar call types and trills with close spectrotemporal features. In particular, larger-sized populations, as expressed by wing length, emitted lower pitched calls. However, the fact that dissimilarities in repertoire and morphology were correlated cannot be explained only by appealing to functional explanations, as the observed intraspecific morphological variability does not seem to be high enough to promote consistent changes in the pool of calls. In this case, historical factors might have contributed to the present pattern of differentiation.     

The relationship between Red-Billed Chough Pyrrhocorax pyrrhocorax (L) breeding populations and grazing pressure on the Calf of Man

A link between the numbers of Loghtan Sheep Ovis aries var. and breeding pairs of Red-Billed Choughs Pyrrhocorax pyrrhocorax has been described for the Calf of Man, Isle of Man, UK, through the analysis of data from 1969 to 1982. Recent research has included further data on populations of Choughs and Sheep grazing, as well as data on brood sizes, Rabbit Oryctolagus cuniculus populations and summer rainfall. Data from 1969 to 1994 show a significant relationship between Sheep and Chough numbers, although data for the period 1983 to 1994 in isolation show no significant association. The mean (± se) brood size ringed in the nest in the period 1959–94 was 2.91 ± 0.011. Between 1972 and 1994, as Sheep numbers peaked, mean brood size decreased, only to rise again as Sheep numbers declined. High Rabbit populations correspond statistically with large Chough broods and increased fledging success. Declines in the Chough breeding population appear to be most closely linked to declines in Rabbit numbers due to outbreaks of myxomatosis, even during periods when Sheep numbers were maintained. The recovery of the Chough population after 1970 corresponds with a change in the effects of myxomatosis outbreaks. Rabbit grazing appears to be equally as important as Sheep grazing in the maintenance of the sward characteristics favoured by breeding Choughs. Combined data on Sheep and Rabbit grazing describe the fluctuations observed in Chough breeding success more fully than data for either grazer alone.     

Factors affecting flock size in the Alpine Chough Pyrrhocorax graculus

The Alpine Chough Pyrrhocorax graculus is a social corvid which now uses food provided by tourist activities in mountain regions (e.g. at ski stations, refuse dumps, picnic areas). Foraging flock size, flock distribution and flock structure of Alpine Choughs were studied in the northern French Alps between 1988 and 1992. Flock size varied during the year, being larger in winter than in summer. Alpine Choughs were not evenly distributed in space, and their numbers depended principally on site-specific factors, such as seasonal human presence. A significant positive correlation existed between Alpine Chough foraging flock size and the local human population. Immature birds gathered in winter in the largest flocks and principally frequented sites with the greatest food availability (e.g. refuse dump). In summer, immatures dispersed, joined small breeding groups and were more homogeneously distributed.     

Effects of behavioural time budgets and nest-building efficiency on male reproductive performance in red bishops ( Euplectes orix )

In this study we have investigated the effect of nest-building behaviour, courtship behaviour, and male–male interactions on male reproductive performance of the red bishop (Euplectes orix), a highly polygynous, colonially breeding weaverbird species. Previous studies on red bishops have revealed that male reproductive success is mainly determined by the number of nests built in a territory, and that reproductive success and the number of nests both vary substantially among males. Here we test whether the high variation observed in reproductive performance is related to male behavioural patterns (as assessed by time-budget analysis) and/or nest-building efficiency (as assessed by the number of fibres incorporated in the nest in unit time). We collected data on male time budgets and analysed behavioural patterns for three breeding seasons. A greater number of nests built was positively related to both the amount of time allocated to nest-building behaviour and the efficiency of nest building. Neither the amount of time spent in courtship behaviour nor the amount of time spent in male–male interactions was related to reproductive success. Male reproductive success, irrespective of the number of nests built, was partly determined by the number of fibres incorporated, suggesting some importance of nest quality in terms of nest chamber density to male reproductive success. There were no trade-offs with regard to time allocated to different behaviour. Instead, males differed with regard to their territory attendance and activity levels, which might be because of differences in energy resources and may thus reflect inherent differences in male body condition and, ultimately, male genetic quality.     

Nest temperature fluctuations in a cavity nester,the southern ground-hornbill

Southern ground-hornbills Bucorvus leadbeateri inhabit savanna and bushveld regions of South Africa. They nest in the austral summer, which coincides with the wet season and hottest daytime temperatures in the region. They are secondary cavity nesters and typically nest in large cavities in trees, cliffs and earth banks, but readily use artificial nest boxes. Southern ground-hornbills are listed as Endangered in South Africa, with reintroductions into suitable areas highlighted as a viable conservation intervention for the species. Nest microclimate, and the possible implications this may have for the breeding biology of southern ground-hornbills, have never been investigated. We used temperature dataloggers to record nest cavity temperature and ambient temperature for one artificial and 11 natural southern ground-hornbill tree cavity nests combined, spanning two breeding seasons. Mean hourly nest temperature, as well as mean minimum and mean maximum nest temperature, differed significantly between southern ground-hornbill nests in both breeding seasons. Mean nest temperature also differed significantly from mean ambient temperature for both seasons. Natural nest cavities provided a buffer against the ambient temperature fluctuations. The artificial nest provided little insulation against temperature extremes, being warmer and cooler than the maximum and minimum local ambient temperatures, respectively. Nest cavity temperature was not found to have an influence on the breeding success of the southern ground-hornbill groups investigated in this study. These results have potentially important implications for southern ground-hornbill conservation and artificial nest design, as they suggest that the birds can tolerate greater nest cavity temperature extremes than previously thought.     

Effects of fleas on nest success of Arctic barnacle geese: experimentally testing the mechanism

Parasites have detrimental effects on their hosts’ fitness. Therefore, behavioural adaptations have evolved to avoid parasites or, when an individual is already in contact with a parasite, prevent or minimize infections. Such anti‐parasite behaviours can be very effective, but can also be costly for the host. Specifically, ectoparasites can elicit strong host anti‐parasite behaviours and interactions between fleas (Siphonaptera) and their hosts are one of the best studied. In altricial bird species, nest fleas can negatively affect both parent and offspring fitness components. However, knowledge on the effects of fleas on precocial bird species is scarce. Research on geese in the Canadian Arctic indicated that fleas have a negative impact on reproductive success. One possible hypothesis is that fleas may affect female incubation behaviour. Breeding females with many fleas in their nest may increase the frequency and/or duration of incubation breaks and could even totally desert their nest. The aim of our study was to 1) determine if a similar negative relationship existed between flea abundance and reproductive success in our study colony of Arctic breeding barnacle geese Branta leucopsis and 2) experimentally quantify if such effects could be explained by a negative effect of nest fleas on female behaviour. We compared host anti‐parasite and incubation behaviour between experimentally flea‐reduced and control nests using wildlife cameras and temperature loggers. We found that flea abundance was negatively associated with hatching success. We found little experimental support, however, for changes in behaviour of the breeding female as a possible mechanism to explain this effect.     

Constrained by available raptor hosts and islands: density-dependent reproductive success in red-breasted geese

In this paper we aim to explain the distribution of red-breasted geese Branta ruficollis over different nesting habitats. To be safe from land predators red-breasted goose colonies were restricted to i) islands on rivers, ii) cliffs with peregrine falcons Falco peregrinus , and iii) the close proximity of snowy owl Nyctea scandiaca and rough-legged buzzard Buteo lagopus nests. Among years nest site availability varied by fluctuations in numbers of owls and buzzards in association with cycles in lemming abundance, but the total number of goose nests found in the study area did not vary. The distribution of geese, in combination with data on reproductive success, suggested a despotic mechanism: at cliffs, goose numbers were constant among years with an invariably high reproductive success, whereas large fluctuations in numbers on islands coincided with opposite trends in success. Apparently, geese nesting with owls or buzzards moved to the few islands present in the study area during years when these birds of prey were absent. Consequently, in such years the average density of geese on islands was more than twice as high as at cliff colonies (5.4 and 2.3 pairs per ha of foraging habitat, respectively). Colony size at cliffs may have been restricted by territorial behaviour of the geese, though there is evidence that, additionally, the host falcons also limited the number of nesting geese. Apparently rare in closely related species, we observed a negative density-dependent effect on reproductive success during the nest phase, and attribute this to limited food resources, reinforced by the high frequency of territorial interactions. This leads to the conclusion that, in addition to predation pressure, nesting density is an important agent in the link between lemming cycles and goose breeding success.     

Competition and coexistence of the European Bee-eater Merops apiaster and the Blue-cheeked Bee-eater Merops persicus in Asia

Studies were conducted over a 10-year period on the supposedly similar European Bee-eater Merops apiaster and Blue-cheeked Bee-eater Merops persicus breeding in mixed and separate colonies in four Asiatic countries. In spring, M. persicus arrived a few days later and laid up to 2 weeks later than M. apiaster. Spatial distributions of the two species were positively associated. They were sympatric and syntopic: more than half of the local breeding ranges overlapped, and many birds bred within sight and sound of the congener. Intraspecific conflict was frequent, but in mixed colonies interspecific conflict was rare. Most M. apiaster nest burrows were dug into cliffs and most M. persicus ones into level ground, but dense mixed colonies occurred only in cliffs. Burrow architecture differed specifically. Diets were qualitatively similar at insect family level but different at the species level, partly because of local variation in availability and partly because of distinct preferences of M. apiaster for small beetles, ants and termites and of M. persicus for large dragonflies and cicadas. Merops persicus was less specialized than M. apiaster and had an airborne insect prey spectrum nearly twice as broad. Diets were more alike where the two birds foraged together than where they foraged separately. There was a high incidence of egg and nestling loss by predation and starvation.
We speculate that each species may prove to breed more successfully in mixed than in monospecific colonies. We propose that the two bee-eaters do not compete for nest sites but may compete for food and coexist unaggressively by trading off food competition against improved breeding success in mixed colonies.     

Nest site selection and efficacy of artificial nests for breeding success of Scarlet Macaws Ara macao macao in lowland Peru

Psittacidae (parrots) have the most threatened species of any bird family in the world. Most parrots are obligate secondary cavity nesters, and can be limited in their breeding success by the availability and quality of nest hollows. However, nesting opportunities for parrots can be increased by provision of artificial nest boxes. The Tambopata Macaw Project has been studying the breeding ecology and natural history of the Scarlet Macaw Ara macao macao in the south-eastern Peruvian Amazon for over 20 years by monitoring natural nest hollows and two types of artificial nest (wooden and PVC). We present data for breeding success in natural and artificial nests over 12 consecutive breeding seasons. The aims of this study were to: (a) determine the nesting requirements and reproductive success of breeding macaws; and, (b) compare the efficacy of the two types of artificial nests and natural nest cavities. Our data showed a high rate of reoccupation of successful nests in consecutive years and that nests in artificial and natural nests had very similar reproductive parameters. Our results indicate that artificial nest types can be used by conservation managers seeking to assist A. macao populations where nest hollows are in short supply, and that artificial nests can contribute important data to natural history studies of species where access to natural nests is limited.     

Diet of the Red-billed Chough Pyrrhocorax pyrrhocorax in south-east Spain

A sample of 140 Red-billed Chough pellets (35 from each season) was collected from communal roosting sites in south-east Spain. In almost every pellet there were three distinct dietary fractions: animal, vegetable and mineral. The animal faction constituted nearly 50% of the pellet volume in each of the four seasons. Wild grains and cultivated cereals were the most important vegetable elements, while 60% of animal prey (n= 3484) were beetles (mainly Tenebrionidae). There was marked seasonal variation in the composition of both the vegetable and animal fractions, the latter associated mainly with variation in the occurrence of Orthoptera, Lepidoptera larvae and Formicidae. Mixed flocks of Choughs and Jackdaws were common in the study area though agonistic interspecific interactions were never observed. The diets of the two species differ in that Choughs eat more beetles and wild seeds, whilst Jackdaws prefer ants and cereals. These observations support the hypothesis that competition from Jackdaws is not responsible for the reduction in the Chough's range.     

Intraspecific brood parasitism: a strategy for floating females in the European starling

In many bird species, there is a floating population of females that are excluded from breeding because of competition for limited breeding resources. Female floaters may enhance their reproductive success by engaging in intraspecific brood parasitism. We studied female floaters in a population of European starlings, Sturnus vulgaris, in order to determine their identity and potential parasitic behaviour. Females were caught after being attracted to nestboxes with artificial nests during 1993-1995. None of the females was known to have a nest of her own at capture but 47% of the females either laid an egg in the nest or carried a fully developed egg within the reproductive tract, indicating that they were intraspecific brood parasites. The floating females were significantly younger and smaller than breeding females. Of 13 females equipped with radiotransmitters and followed daily, all but one started a breeding attempt of their own after 3-8 days and the majority settled as secondary females or mated with males where the original female had disappeared. This suggests that females that are unable to compete successfully for nest sites or males early in the breeding season may use intraspecific brood parasitism to enhance reproductive success during the period that they are constrained from breeding. The importance of settling rapidly because of a seasonal decline in reproductive success may also promote the evolution of intraspecific brood parasitism in the starling. The relative reproductive success of combining egg dumping with breeding compared with traditional breeding will depend on the costs of delaying breeding as well as the probability of finding a mate later in the breeding season. Copyright 1999 The Association for the Study of Animal Behaviour.     

Factors affecting nesting success of a bird assembly in the central Monte Desert, Argentina

We estimated nesting success of a breeding bird assemblage in the central Monte Desert in four breeding seasons (1995–1999). We analyzed the effect of year, period within the breeding season, nest type, and nest location on daily survival rates (DSR). Averaged nesting success of passeriforms in natural habitats was very low (<20% on average, n=12 species). DSR tended to be lower during the egg-laying stage in Furnariidae, Rhinocryptidae, Mimidae, and Emberizidae. Tyrannidae had the highest DSR for all nesting stages and Emberizidae the lowest. Predation was the main cause of nest failure (>90% on average, n=12 species). DSR differed among years, being higher in the driest year (1995) and lower in the most humid year (1997). Nests initiated in the first half of the breeding season had higher DSR than those initiated in the second half. Closed nests had similar survival rates to open nests. DSR differed according to species of plant that supported the nest. Nest predation in the central Monte Desert may be an important selective pressure in birds' life history, which is in accordance with other results for South American temperate birds.     

Ants as a Major Food Source of the Chough

Few birds are more captivating in their behaviour, more exciting in their aerial manoeuvres, than the Chough, a species now restricted to the wilder parts of our western coasts. This field study of its specialised feeding habits shows how closely its future welfare may depend upon the continuance of a grazing regime on islands and headlands within its range.     

The diet of a declining Red-billed Chough Pyrrhocorax pyrrhocorax population on Islay,Scotland

Capsule: We quantify Red-billed Chough Pyrrhocorax pyrrhocorax diet from faecal samples collected on Islay. Dung invertebrates formed the majority of prey biomass in dune pasture and Tipulid larvae in mixed pasture but Aphodius larvae were scarce in the diet, whereas they had been a major component in the 1980s. There are management implications from the indication of a reduction in availability of preferred food.     

Rock sparrow song reflects male age and reproductive success

The evolution of mating signals is closely linked to sexual selection. Acoustic ornaments are often used as secondary sexual traits that signal the quality of the signaller. Here we show that song performance reflects age and reproductive success in the rock sparrow (Petronia petronia). In an Alpine population in south-east France, we recorded the songs of males and assessed their genetic breeding success by microsatellite analysis. In addition to temporal and spectral song features, we also analysed for the first time whether the sound pressure level of bird song reflects reproductive success. Males with higher breeding success sang at a lower rate and with a higher maximum frequency. We found also that older males gained more extra-pair young and had a higher overall breeding success, although they also differed almost significantly by having a higher loss of paternity in their own nests. Older males could be distinguished from yearlings by singing at lower rate and higher amplitudes. Our findings suggest that song rate may be used as a signal of age and together with song pitch as a signal of reproductive success in this species. Alternatively, younger and less successful males might try to compensate their inferior status by increased song rates and lower pitch. Independent of age and quality, high-amplitude songs correlated with paternity loss in the own nest, suggesting that in this species song amplitude is not an indicator of male quality but high-intensity songs may be rather a response to unfaithful social mates.