Factors affecting the availability of invertebrate food for the chough, Pyrrhocorax pyrrhocorax L
Most of the fieldwork for this study was conducted on the island of Islay, in the Inner Hebrides, the stronghold of the chough, Pyrrhocorax pyrrhocorax L., in Scotland. The aims of this study were to provide baseline data on the phenology of potential invertebrate foods of the chough, and to provide a greater understanding of the factors affecting these invertebrate populations. The literature concerning (a) the chough in Britain and its feeding ecology, (b) the invertebrate fauna of pasture, (c) the invertebrates associated with cow dung, (d) ivermectin and its effect on the invertebrate fauna associated with cattle dung, and (e) the multivariate analysis methods used in this study, is reviewed. An area of heather moorland and four pastures were selected on Islay. Invertebrates were collected from these sites between January 1988 and November 1989 using pitfall traps, and by sampling soil and cow pats. The data obtained was analysed using two multivariate analysis methods -Two-Way-Indicator-Species- Analysis (TWINSPAN) and Detrended Correspondence Analysis (DECORANA). Information on 62 surface-active taxa was obtained from pitfall trapping. Although seasonal taxa assemblages were recognized, the distribution of the invertebrate communities was primarily related to soil moisture content. Grazing intensity and seasonality were also important factors determining the composition of the invertebrate fauna at each site. The taxa active during the summer and winter at the two sand grassland sites, did not appear to differ as markedly as at the other sites sampled. Figures showing the seasonal activity of some of the frequently occurring taxa at each site considered potential chough prey items are provided. Surface-active potential chough food items were present, at all the sites investigated, throughout the year. Soil-sampling provided information on 34 taxa. As with the surface-active fauna, the primary factor influencing the soil fauna was soil moisture content. The time of year was also an important factor governing the soil fauna composition, with the majority of taxa occurring in low numbers during the summer months at all the sites sampled. Figures indicating the seasonal occurrences of some of the taxa considered potential chough prey items at each site are provided. Soil did not appear to be a good source of potential prey items for the chough during the summer months, although, as a result of seasonal increases in size, certain taxa, e.g. Tipulidae larvae, may have been more 'worthwhile' prey items at this time of year than at any other. Information on 54 taxa was obtained from sampling cow dung. Seasonality and age of the dung were very important in determining the composition of the dung fauna. The seasonal variations in the fauna associated with the cow pats are described. Potential chough prey items were associated with cow dung, in any stage of decay, throughout most of the year. Only during the period from October/November to January did there appear to be a lack of suitably sized potential prey items in the dung. The 'summer' months, when fresh dung contained large numbers of beetle adults and developing fly larvae, and late autumn, when pats deposited during the summer months are old enough for the large numbers of Aphodius spp. larvae present to have attained a reasonable size, were considered to be the times at which cow dung presented the best feeding opportunities for the chough. Fifty taxa were identified in samples of chough faeces. Multivariate analysis of these data indicated that the seasonal availability of prey items was the most important factor influencing chough diet throughout the year. Soil-dwelling Tipulidae (January to July) and Bibionidae (January to April) larvae, dung-associated insects (during the spring, and late summer and autumn), and surface-active insects (during the summer) were important invertebrate components of the diet. Cereal grains were extremely important supplementary food items during the early winter months, when invertebrate availability was low. An experiment was conducted at the College to investigate the effects on the dung fauna of spiking cow dung with 2.0, 1.0, or 0.5 mg/kg dung of ivermectin. Pats were placed on pasture between May and September 1988. The pats were lifted, and the soil beneath sampled, after 15 to 90 days exposure. A total of 65 taxa were identified. These data were analysed using TWINSPAN, DECORANA and Canonical Correspondence Analysis (CANOCO). The major factors determining the invertebrate fauna of the pats were length of exposure, exposure date, and ivermectin presence/absence. Ivermectin markedly affected the fauna associated with the pats. Pats exposed in June and August degraded faster than those exposed in May or September. In June, the ivermectin-treated pats degraded significantly slower than the control pats. An attempt to extract ivermectin from cow dung for analysis by high-performance liquid chromatography is described. This proved unsuccessful and the reasons for this failure, and possible improvements, are discussed. The main conclusions of this study are: (1) that Tipulidae larvae are extremely important components of the chough's diet on Islay, and that the climatic conditions of the island favour these insects; (2) livestock farming on Islay, especially the out-wintering of cattle, provides essential feeding opportunities for the chough, as, (a) gazing animals produce the short sward preferred by the chough as a feeding habitat, (b) large numbers of insects are associated with the dung of these animals, and (c) supplementary feed provided for the cattle in winter also provides an essential alternative food source for the chough at a critical time; (3) the chough's preference on Islay for feeding in sandy, coastal pasture is due to the fact that these sites, (a) contain a variety of suitable invertebrate prey items throughout most of the year, (b) are normally intensively grazed and so contain large amounts of dung with its associated fauna, and (c) are used for out-wintering cattle and therefore cereal grains can be found there; (4) treating cattle with ivermectin could have an adverse effect on the chough as it reduces the number and variety of invertebrates associated with the dung, an important food source for the birds, especially in spring and autumn.