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.     

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.     

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.     

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.     

A comparative analysis of the breeding biology of the Red-billed Chough Pynhocorax pyrrhocorax and the Alpine Chough P. graculus coexisting in the Alps

We compare the breeding biology of a small population of Red-billed Chough Pyrrhocorax pynhocorax with that of the Alpine Chough P. graculus coexisting in the western Italian Alps. The analysis focused on nest site choice, timing of breeding, reproductive success and parental effort (assessed by means of behavioural indices such as food provisioning rates and time budgets inside and close to the nest). The Alpine Chough nested in a greater variety of sites: natural cliffs, windows of a dam, pot-holes and abandoned buildings; the Red-billed Chough used only natural cliffs. Laying dates also differed significantly, with the Red-billed Chough breeding one month earlier than its congener. Differences in the feeding ecology of those species might be responsible for the observed temporal partitioning, thus facilitating coexistence. Breeding success and reproductive behaviour were similar, and the species differed in only one out of eight breeding parameters. It is suggested that similar strong constraints on breeding behaviour have resulted in the similarities in behaviour seen in these species.     

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.     

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.     

Social structuring at a communal roost of Choughs Pyrrhocorax pyrrhocorax

The social structure of a communal roost of Choughs Pyrrhocorax pyrrhocorax was studied between July and September 1985. Different age classes within the roost were spatially segregated. Third-year birds were significantly more likely to attack another Chough and roosted in the densest part of the roosting flock. First-year birds roosted significantly lower down the roost cliff and on the periphery of the flock.     

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.     

Demographic consequences of prey availability and diet of Red-billed Choughs Pyrrhocorax pyrrhocorax

Capsule Variation in prey availability appears to influence Chough fledging success and juvenile survival.

Aims To determine seasonal and annual variations in Chough prey and how these influence fledging success and juvenile survival.

Methods Chough faeces (n = 437, 3905 invertebrates) were collected year-round and analysed to determine diet composition. Seasonal and annual variation in prey abundance in Chough foraging habitat was assessed using pitfall trapping (n = 747, 27 124 invertebrates) between 1996 and 2003. Fledging success was estimated for a population of 12 breeding pairs; juvenile survival was estimated by year-round resighting (n < 2500) of individual birds that were colour-ringed as nestlings.

Results From April to June a high biomass of prey and a great variety of species were observed. Between July and October, both biomass and species diversity were reduced; prey consisted mainly of ants and a few beetle species. From November to March, biomass availability was intermediate, with the diet consisting mostly of Tipulidae larvae, plants and dung beetles. Chough juvenile monthly survival was low in August, November and December. Annual variation in fledging success was correlated with prey biomass availability in May.

Conclusion Seasonal variation in the availability of prey species and their biomass influences Chough demography.     

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.     

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.     

The influence of flocking on the foraging behaviour of the chough (Pyrrhocorax pyrrhocorax) and the Alpine chough (P. graculus) coexisting in the Alps

We studied the flocking and foraging behaviour of the chough Pyrrhocorax pyrrhocorax and the Alpine chough P. gruculus coexisting in the south-western Italian Alps in order to evaluate the costs and benefits of foraging in single- and mixed-species flocks.
In the single-species context, flock size significantly affected the foraging behaviour of the Alpine chough; in larger flocks, the birds stayed for a shorter time in a patch and fed more quickly than in smaller flocks. Flock size did not significantly affect the foraging behaviour of the chough, probably because of the small number of individuals per flock.
The propensity for mixed-species flocking was rather low. The observed frequencies of single-species flocks of choughs and Alpine choughs were significantly higher than those expected on the basis of random flocking, whereas the observed frequencies of mixed-species flocks of the two species were lower than those expected. The stay times became significantly shorter for the chough in the presence of the Alpine chough. Moreover, feeding rates of the Alpine chough were significantly lower in the presence of the dominant chough.
The present study does not confirm the hypothetical foraging advantages of flocking. In single-species flocks, the benefits for the Alpine chough (higher feeding rates in larger flocks) were roughly compensated by the costs (shorter stay times in larger flocks), whereas the chough apparently neither gained benefits nor endured costs.
In mixed-species flocks, the Alpine chough sustained costs due to a reduction of feeding rates and the chough suffered costs due to a reduction of stay times. Hence, on average, single-species flocking gives no evident foraging advantages to either the chough or the Alpine chough, whereas mixed-species flocking provides some disadvantages for both species.     

The food of the Common frog, Rana temporaria, on high moorland in northern England

Analysis of the stomach contents of 427 Common frogs, Rana temporaria L., captured on an area of moorland typical of the northern Pennines, showed that this species is an unspecialized and opportunistic feeder in this habitat.
There was a distinct temporal variation in the food eaten which was closely related to its availability. Adult Tipulidae and Lepidoptera larvae each formed a fifth of the diet by volume, while Phalangidae, Coleoptera, and Gastropoda, each formed about a tenth. The diet of the frogs changes as they grow. Small frogs fed almost entirely on small items. The bigger the frog the wider the range of items eaten and the larger the items which were predominantly eaten.     

Captive Husbandry and Socialization of the Red‐Billed Chough (Pyrrhocorax pyrrhocorax)

Since the 1970s, Paradise Park Wildlife Sanctuary in Cornwall, United Kingdom, has built up a captive flock of red‐billed choughs Pyrrhocorax pyrrhocorax and over 30 years has developed successful methods of keeping, breeding, and appropriately socializing them in captivity. A total of 77 nests reached the egg stage with 27 nests producing at least one young and 48 young fledging in total. Several components are important in achieving successful breeding and socialization. Provision of live food, especially ant's eggs and small mealworms and crickets, in the first days after hatching is essential, improving the condition of adults and survival of nestlings. Situating aviaries in quiet areas, away from public view, is important. Socialization in family groups during the winter months and allowing choughs to choose partners induces better compatibility. Introduction of nest cameras greatly improved young survival through early identification of health problems enabling treatment of young between hatching and 10‐days old, when mortality is otherwise highest, and enabling precautionary medication shortly after hatching. We show that clutch size increases significantly with female age and that direct intervention such as artificial egg incubation and hand rearing can be successful and worthwhile, but its requirement is reduced by closer monitoring. Red‐billed choughs provide a good model species to further develop captive management and release techniques that can then be applied to critically endangered species that show similar social and long‐learning behaviors. Captive breeding programs can play an important role in such work through provision of suitable birds and supporting avicultural expertise. Zoo Biol. 31:725‐735, 2012. © 2012 Wiley Periodicals, Inc.     

Summer food of the golden plover Pluvialis apricaria at Hardangervidda, southern Norway

Golden plover stomachs were examined through the breeding season al Hardangervidda, southern Norway. The main foods were Diptera larvae (in particular Tipula excisa ), adult Coleoptera (notably Otiorrhynchus dubius and Carabidae), and in May and August berries of Empetrum hermaphroditum. Also some larvae of Coleoptera and Lepidoptera were taken. Adult Coleoptera were considerably more common in the diet than expected from abundance data, while Araneae were unimportant as food in spite of a high abundance. The males take the major parental duties after hatching, allowing the females a more undisturbed feeding, supposed to be strongly advantageous for them due to close proximity of egg-laying and the start of the postnuptial moult. This is reflected in the diet: in July males showed a larger tropic diversity, and ate more adult insects and less insect larvae than the females.     

Similarity and specialization of the larval versus adult diet of European butterflies and moths

Many herbivorous insects feed on plant tissues as larvae but use other resources as adults. Adult nectar feeding is an important component of the diet of many adult herbivores, but few studies have compared adult and larval feeding for broad groups of insects. We compiled a data set of larval host use and adult nectar sources for 995 butterfly and moth species (Lepidoptera) in central Europe. Using a phylogenetic generalized least squares approach, we found that those Lepidoptera that fed on a wide range of plant species as larvae were also nectar feeding on a wide range of plant species as adults. Lepidoptera that lack functional mouthparts as adults used more plant species as larval hosts, on average, than did Lepidoptera with adult mouthparts. We found that 54% of Lepidoptera include their larval host as a nectar source. By creating null models that described the similarity between larval and adult nectar sources, we furthermore showed that Lepidoptera nectar feed on their larval host more than would be expected if they fed at random on available nectar sources. Despite nutritional differences between plant tissue and nectar, we show that there are similarities between adult and larval feeding in Lepidoptera. This suggests that either behavioral or digestive constraints are retained throughout the life cycle of holometabolous herbivores, which affects host breadth and identity.     

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.     

Traditional farming and key foraging habitats for chough Pyrrhocorax pyrrhocorax conservation in a Spanish pseudosteppe landscape

1. Agricultural land devoted to low-intensity practices supports many declining bird species in Europe. The potential effects of intensification or abandonment of traditional farming practices are assessed from the point of view of the conservation of the chough Pyrrhocorax pyrrhocorax .
2. The study was conducted in Los Monegros (north-east Spain), in a pseudosteppe landscape of special relevance for this species. In this area traditional farming survives in the form of low-intensity arable systems including fallowing and extensive sheep grazing.
3. Foraging choughs showed a strong selection for field margins, avoided any remnant of the original scrubland and halophitic vegetation, and utilized other habitats maintained by traditional crop rotation (e. g. fallow, stubble) at different times of the year.
4. A highly complex pattern of habitat selection was found which was not only related to seasonal changes in habitat availability, but also differed between territorial pairs and the non-breeding population. This emphasizes the importance of taking all the different components of bird populations into account when analysing interactions between habitat selection and the annual farming cycle.
5. Chough habitat selection suggests that both agricultural intensification and land abandonment would have detrimental consequences for this species in Los Monegros. The maintenance of traditional farming is recommended, especially by promoting the use of long fallow rotations grazed extensively by livestock, encouraging the increase of grassland edges around landholdings and sand roads, and maintaining a low input of biocides. These management measurements could also favour most other endangered species of steppe birds, and could be supported by the establishment of Zonal Programmes under the CAP Agri-Environment Regulation (2078/92).     

Habitat selection and diet of the Capercaillie Tetrao urogallus in Abernethy Forest,Strathspey, Scotland

Capsule Woodland characteristics are important for fully grown birds and for broods.

Aims To describe habitat selection and diet of the Capercaillie in ancient native pinewoods and Scots Pine Pinus sylvestris plantations, in order to provide a guide for forest management.

Methods Tree selection was studied in winter by locating droppings under trees along transects, and comparing those trees used by Capercaillie with those not used. Habitat selection by broods was studied by following radiotagged chicks or females with broods. Diet was described from the analysis of droppings.

Results In winter, Capercaillie preferred Scots Pines with wide crowns and large branches, and little surrounding Heather Calluna vulgaris, but avoided forest tracks. Female Capercaillie occurred in stands with a higher density of pines compared to males. The diet of fully grown birds comprised primarily Scots Pine needles between October and April. Scots Pine pollen cones were eaten during May, June and July. Bilberry Vaccinium myrtillus leaves and fruit were eaten in summer and autumn. Habitats used by broods had a higher percentage of Vaccinium sp. and smaller Scots Pines compared with random sites. Chicks hatch at the beginning of June, and in their first two weeks, their diet comprised Formicidae, Lepidoptera larvae and Coleoptera. In July and August, Lepidoptera larvae were rarely eaten but a higher percentage of chicks took vegetable material (particularly Bilberry fruit). In June, smaller numbers of invertebrates were generally trapped in mires of wooded bogs compared with areas of Heather and Vaccinium sp. in old woodland. Areas of Vaccinium sp. had the highest numbers of Formicidae, Coleoptera and Diptera, and especially Lepidoptera larvae. Numbers of Lepidoptera larvae declined during June and July as larvae pupated, explaining the reduction of Lepidoptera larvae in the diet of older chicks.

Conclusions Habitat management for Capercaillie should aim to provide old Scots Pine forest with a Vaccinium-rich field layer. Within Scotland, this may entail increasing the tree density in ancient native pinewoods but decreasing it in pine plantations. The removal or closure of tracks might increase the amount of woodland available to Capercaillie, by reducing human disturbance.