Prey abundance and the strength of interference in a foraging shorebird

Interference is an important component of food competition but is often difficult to detect and measure in natural animal populations. Although interference has been shown to occur between oystercatchers Haematopus ostralegus L. feeding on mussels Mytilus edulis L., four previous studies have not detected interference between oystercatchers feeding on cockles Cerastoderma edule L. In contrast, this study detected interference between cockle-feeding oystercatchers in the Baie de Somme, France. Prey stealing (kleptoparasitism), one of the main causes of interference between mussel-feeders, also occurred between oystercatchers in the Baie de Somme. The kleptoparasitism rate was related to the natural variation in the food supply, tending to be higher when cockles were rare. Feeding rate was negatively related to competitor density, so providing evidence for interference, but, as in mussel-feeders, only above a threshold density of about 50–100 birds ha⁻¹. The strength of interference at a fixed competitor density was related to the cockle food supply, usually being greater when cockles were rare. Previous studies probably failed to detect interference between cockle-feeders because competitor densities were too low, or cockles were too abundant, or because they were not conducted during late winter when interference is most intense. The study shows that natural variation in the food supply can influence the strength of interference within an animal population and provides support for those behaviour-based interference models which predict that the strength of interference will be greatest when competitor densities are high and prey scarce.     

The mechanisms of interference competition: two experiments on foraging waders

Models of population dynamics that include interference competitionhave often been applied to foraging waders and less so to otherforagers, even though these models are, in principle, generallyapplicable. At present, however, it is still unclear whetherinterference competition is of importance for foraging waders.To support this idea experimental evidence and knowledge ofthe mechanisms underlying interference effects are required.We experimentally determined the relationship between foragerdensity and foraging success in two wader species: the red knot(Calidris canutus) and the ruddy turnstone (Arenaria interpres).With each of the two species, we conducted an experiment consistingof 300 one-min trials. In these trials we scored the behaviorand the foraging success of focal individuals at specific combinationsof bird and prey density. Irrespective of prey density, individualsof both species discovered fewer prey items at higher bird densities.Despite this, only in turnstones did intake rates decline withincreasing bird density. Knots compensated for a lower prey-discoveryrate by rejecting fewer prey items at higher bird densities.In knots, bird density had a complex, nonmonotonic effect onthe time spent vigilant and searching. In turnstones the maineffect of increased bird density was a reduction in the prey-encounterrate, that is, the reward per unit search time. Effects on thetime spent vigilant and the time spent searching were less pronouncedthan in knots. Thus, the mechanistic basis of the effects ofbird density was complex for each of the two species and differedbetween them.     

Density dependence and the stabilization of animal numbers

Summary Latto and Hassell (1987) disagree with the conclusion of Den Boer (1986), that the winter moth population at Wytham Wood, studied by Varley and Gradwell, was not regulated. They attempt to demonstrate regulation by means of a simulation model. In the present paper the validity of this model is tested step by step. The fixing of the initial and final densities, as practised by Den Boer and rejected by Latto and Hassell, did not prevent population explosions and extinctions, as was assumed by Latto and Hassell. It is shown that the deterministic formulation of the density dependence of pupal predation, as used by Latto and Hassell, deviates systematically from the field data. Replacing the values of the key-factor (k₁) by random values drawn from a normal distribution (Latto and Hassell) affects the dynamics such that the ability of pupal predation to govern density is improved in the model. Changing mortalities other than the key-factor does not significantly influence the pattern of fluctuations nor the limits of density. Models should leave intact the essentials of the reality under study, while removing distracting elements (Levins 1968). As both the timing of the key-factor, and its correlation with pupal predation are essential features of the winter moth population at Wytham Wood between 1950 and 1968, the model of Latto and Hassell does not apply to this population. By simply changing log₁₀ (eggs/female) it is shown that the power of the density dependence of pupal predation to govern possible trends in density of the winter moth population at Wytham Wood is weak. On the other hand, the model of Latto and Hassell gives insight into the conditions that might favour regulation of numbers. Although the model of Poethke and Kirchberg (1987) preserves more features of the pertinent winter moth population than that of Latto and Hassell (1987) it still deviates in one essential aspect: the succession in time of both the (coupled) mortalities and the deviations from the deterministic density dependence are taken at random. Therefore, also this model is still too far from the field population to be a sound base for the statistical speculation proposed by Poethke and Kirchberg.Communication No. 339 of the Biological Station, Wijster     

Does landscape structure affect resource tracking by avian frugivores in a fragmented Afrotropical forest?

Frugivorous species heavily depend on patchy food resources and are believed to track these in space and time, thereby providing an important seed dispersal function that might be critical toward the regeneration of fruiting plants. However, isolation of suitable food patches due to habitat fragmentation or changes in landscape connectivity may hamper food tracking behaviour and adversely affect populations of both frugivores (through starvation) and food plants (through interruption of seed dispersal). We here test whether density fluctuations in four frugivorous Afrotropical bird species were larger and/or matched fluctuations in ripe fruit densities better in study plots embedded in large tracts of indigenous forest than in equally-sized plots embedded in cultivated lands. We compared these results with those of four non-frugivorous species (out-group) which were not expected to track fruit resources. Whereas densities of both frugivores and fruit crops strongly fluctuated in space and time, these fluctuations were not synchronised, nor did the level of synchrony differ in relation to matrix type. For some but not all bird species, lower densities and smaller temporal fluctuations in forest plots surrounded by cultivation may reflect decreased mobility. The observed fluctuations in bird densities most likely reflect exchange with the surrounding landscape matrix, suggesting that small pockets of fruiting trees in farmland may comprise critical food resources for frugivores inhabiting highly fragmented landscapes, apart from increasing connectivity for both bird and seed dispersal.     

Predator interference emerging from trophotaxis in predator–prey systems: An individual-based approach

An individual-based model describing predator–prey interactions within a closed rectangular habitat was developed to study how different assumptions about the individual movements lead to the emergence at the population level of various kinds of prey- and predator-dependence in the spatially aggregated trophic function.In addition to random walk, both species are capable of directional movement, i.e., the model accounts for the predator prey-taxis and evasion of predators by prey individuals. The taxis stimulus of each species is the odour of the other species, which is distributed continuously in space. Spatial behaviour of individuals is determined by the specific response to the odour gradient and the tendency to maintain the taxis velocity.In order to facilitate the assessment of the trophic function, the model allows removing the effect of demographic density variations on the predator ration, keeping population sizes constant.Analyzing the dependence of the trophic function with the average predator density, we found that, depending on the intensity of taxis, the predator population exhibits various degrees of interference, from very low to very high values. In particular, a moderate taxis generates distinct levels of interference including the ratio-dependent case. The letter maximizes the average consumption rate.A new generalized function containing ratio-dependence and prey-dependence as special cases, at high and low population abundances, is suggested. This trophic function fits the simulated data better than the Hassell–Varley–Holling expression does.     

Experimental functional response and inter-individual variation in foraging rate of teal (Anas crecca)

The functional response, i.e. the change in per capita food intake rate per time unit with changed food availability, is a widely used tool for understanding the ecology and behaviour of animals. However, waterfowl remain poorly explored in this context. In an aviary experiment we derived a functional response curve for teal (Anas crecca) foraging on rice (Oryza sativa) seeds. We found a linear relationship between intake rate and seed density, as expected for a filter-feeder. At high seed densities we found a threshold, above which intake rate still increased linearly but with a lower slope, possibly reflecting a switch from filter-feeding to a scooping foraging mode. The present study shows that food intake rate in teal is linearly related to food availability within the range of naturally occurring seed densities, a finding with major implications for management and conservation of wetland habitats.     

Sex-Related Differences in the Trade-Off between Foraging and Vigilance in a Granivorous Forager

The relationship between intake rate and food density can provide the foundation for models that predict the spatiotemporal distribution of organisms across a range of resource densities. The functional response, describing the relationship between resource density and intake rate is often interpreted mechanistically as the relationships between times spend searching and handling. While several functional response models incorporate anti-predator vigilance (defined here as an interruption of feeding or some other activity to visually scan the environment, directed mainly towards detecting potential predators), the impacts of environmental factors influencing directly anti-predator vigilance remains unclear. We examined the combined effects of different scenarios of predation risk and food density on time allocation between foraging and anti-predator vigilance in a granivorous species. We experimentally exposed Skylarks to various cover heights and seed densities, and measured individual time budget and pecking and intake rates. Our results indicated that time devoted to different activities varied as a function of both seed density and cover height. Foraging time increased with seed density for all cover heights. Conversely, an increased cover height resulted in a decreased foraging time. Contrary to males, the decreased proportion of time spent foraging did not translate into a foraging disadvantage for females. When vegetation height was higher, females maintained similar pecking and intake rates compared to intermediate levels, while males consistently decreased their energy gain. This difference in anti-predator responses suggests a sexually mediated strategy in the food-safety trade-off: when resource density is high a females would adopt a camouflage strategy while an escape strategy would be adopted by males. In other words, males would leave risky-areas, whereas females would stay when resource density is high. Our results suggest that increased predation risk might generate sexually mediated behavioural responses that functional response models should perhaps better consider in the future.     

Balancing food and density‐dependence in the spatial distribution of an interference‐prone forager

Foraging distributions are thought to be density‐dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density‐dependent and density‐independent effects on the spatio‐temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density‐independent interactions with the food landscape. Spatial distributions were quantified using high‐resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour‐based optimality models against these tracking data. Density‐independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference‐prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference‐free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density‐independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density‐independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter‐individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised.     

The effect of multiple parasitoid introductions upon the equilibrium value of host density

Summary Four models of host-parasitoid population variation are examined to study the effect of additional parasitoid introductions. Only one of the models predicts that the equilibrium value of host population increases with such introductions. The other three, that of Holling, of Nicholson, and its recent modification by Hassell and Varley, predict a fall in the equilibrium of host numbers. Thus it would appear that the practice of multiple introductions in biological control is correct when the host-parasitoid relationship can be modelled by these latter systems.     

Prediction of bird-day carrying capacity on a staging site: a test of depletion models

1. The carrying capacity of a site for migratory water birds, expressed in bird-days, can be of particular conservation value. Several attempts have been made to model this carrying capacity using ideal free distribution models such as, for instance, depletion models, in which the distribution is fully determined by exploitative competition. 2. In the tests of depletion models carried out so far, no alternative models were compared; rather, one specific model was tested. We tested whether bird-days were more in accordance with birds depleting the food resource (a1) until a critical food density which just enabled survival or (a2) until a threshold food density which renders the site as profitable as an alternative site; and birds (b1) satisfying their daily requirements or (b2) maximizing daily intake. 3. We studied Bewick's swans feeding on below-ground tubers of fennel pondweed in one part of an autumn staging site. In most years between 1995 and 2005, we measured tuber biomass densities around September, November and March, and counted swans daily during their stopover in October. 4. The best fit between observed and predicted bird-days was obtained by assuming that the swans were maximizing their daily intake and depleting the tubers until a threshold biomass density (which yielded the same energetic return as the alternative food source after accounting for a small part of the initial tuber biomass being out of reach of the swans). Also in line with daily intake maximization, the daily feeding time did not differ from 10 h day(-1), the value predicted for Bewick's swans based on their feeding costs. 5. Our results suggests that the applicable model to calculate carrying capacity may depend strongly on whether birds use a site to stopover or to winter, because it determines whether the birds are more likely to use a threshold or critical food density, and to behave as energy maximizers or satisficers.     

Biomechanics and foraging profitability: an approach to assessing trophic needs and impacts of diving ducks

A biomechanical model of underwater locomotion is described, and data required by the model presented for 3 species of diving duck (Aythya spp.). Based on field observations of behavior and foods consumed, the model is used to estimate energy costs of foraging and minimum food intake rates of canvasbacks (Aythya valisineria) in two habitats in North Carolina. Increased water depth from 0.5 m in Lake Mattamuskeet to 1.5 m in Pamlico Sound increased the net cost of time spent foraging at the bottom by 43%. Biomechanical calculations are combined with data on intake rates at different food densities (Takekawa, 1987) to determine minimum food densities for profitable foraging in Lake Mattamuskeet. Field observations of behavior are used to adjust minimum intake per dive for unsuccessful dives spent locating food patches. Density and dispersion of plant tuber foods in Lake Mattamuskeet, before and after the fall staging period, suggest that the fraction of habitat with tuber densities above a profitability threshold is more critical to canvasbacks than average tuber density. Such factors are important in relating bird energy requirements and benthic sampling data to carrying capacity and total area of usable habitat. The proportion of foods that can be fed upon profitably also determines the fraction of food organisms subject to depletion as components of trophic pathways.     

A model describing chemical interference caused by decomposing residues at different densities of growing plants

In literature, the biological response of plants to phytochemicals has been modelled and then used to simulate phytotoxicity caused by plant residues during decomposition. According to the resulting residue allelopathy model, stimulation dominates in the beginning of the residue decomposition process for a short while. Thereafter, severe inhibition is predicted to occur rapidly, until stimulation gradually re-emerges at the later stages of residue decomposition. Also in literature, direct chemical interference has been shown to be density-dependent; with increasing target plant density, the effects of phytochemicals are diluted. As a result, inhibition is the most probable outcome in density-dependent phytochemical interactions at low target plant densities, but phytotoxic effects often become stimulatory as target plant density increases. In this paper, these models that have been reported in literature are combined. While the original residue allelopathy model predicts inhibitory effects in most cases, the new density-dependent extension of the residue allelopathy model predicts that the density of target plants determines whether or not inhibition occurs. According to the new model, the intensity of inhibition decreases and the final stimulatory period begins earlier if target plant density increases. Therefore, combining the effects of density-dependency to the residue allelopathy model enhances our understanding of chemical interference. In addition, the new model may partially explain why several field studies have not observed chemically driven inhibitory effects similar to those observed in laboratory experiments.     

Modelling irruptions and population dynamics of the great spotted woodpecker – joint effects of density and cone crops

Irruptive migrants are partially migrating species showing pronounced temporal variation in the number of migrants. Occasional irruptions are often explained by increased population density (per area) or lack of food. Similarly to population dynamics, these explanations are not mutually exclusive, but probably act in concert. Here we do a parallel analysis on the migration intensity and population dynamics of an irruptive migratory bird species, the great spotted woodpecker Dendrocopos major, in southern Finland. Both analyses include simultaneous effects of breeding density and cone crops of Scots pine Pinus sylvestris and Norway spruce Picea abies. A novel model for statistical analyses of irruptive migration intensity is developed. Population dynamics are investigated using log‐linear state‐space models. Woodpecker migration intensity is well explained by lack of spruce cones and increases proportionally to population density. Population dynamics is most successfully described with a density dependent model, where the natural logarithm of spruce cone availability previous autumn has a strong positive effect on population growth. This is likely to be due to lowered winter mortality and lowered emigration rates resulting from plentiful food availability. Although the species uses pine cones as a food resource in winter, the estimated impacts of pine cone crops on migration intensity and population dynamics are low, probably due to low annual variation in crop comparing to spruce cones. Large irruptions also tend to occur earlier in season, often before cones are a crucial food resource for the woodpeckers, suggesting that the woodpeckers use the amount of maturing cones as a cue for upcoming conditions. Our results show that the ecological role of different resources can differ considerably from the expected pattern based on the proportion of the resources in a species’ diet. Advantages of examining patterns of bird migration, population density and dynamics in unison are apparent.     

Species-specific features affect the ability of census-derived models to map winter avian distribution

We identify autoecological traits of bird species that influence the accuracy of predictive models of species distribution based on census data obtained from stratified sampling. These models would serve as a complementary approach to the development of regional bird atlases. We model the winter bird abundance of 64 terrestrial bird species in 77 census plots in Central Spain (Madrid province), using regression tree analyses. The predicted distribution of species density derived from statistical models (birds/10 ha) was compared with the published relative abundances depicted by a very accurate regional atlas of wintering birds (birds observed per 10 h). Statistical models explained an average of 41.7% of the original deviance observed in the local bird distribution (range 19.6–79.3%). Significant associations between observed relative abundances (atlas data) and predicted average densities in 1×1 km squares within 10×10 km UTMs were attained for 44 out of 64 species. Interspecific discrepancies between predicted and observed distribution maps decreased with between-year constancy in regional bird distribution and the degree of ecological specialization of species. Therefore, statistical modeling using census localities allowed us to depict geographical variations in bird abundance that were similar to those in the quantitative atlas maps. Nevertheless, bird distributions derived from statistical models are less reproducible in some species than in others, depending on their autoecological traits.     

Linking foraging decisions to residential yard bird composition

Urban bird communities have higher densities but lower diversity compared with wildlands. However, recent studies show that residential urban yards with native plantings have higher native bird diversity compared with yards with exotic vegetation. Here we tested whether landscape designs also affect bird foraging behavior. We estimated foraging decisions by measuring the giving-up densities (GUD; amount of food resources remaining when the final forager quits foraging on an artificial food patch, i.e seed trays) in residential yards in Phoenix, AZ, USA. We assessed how two yard designs (mesic: lush, exotic vegetation; xeric: drought-tolerant and native vegetation) differed in foraging costs. Further, we developed a statistical model to calculate GUDs for every species visiting the seed tray. Birds foraging in mesic yards depleted seed trays to a lower level (i.e. had lower GUDs) compared to birds foraging in xeric yards. After accounting for bird densities, the lower GUDs in mesic yards appeared largely driven by invasive and synanthropic species. Furthermore, behavioral responses of individual species were affected by yard design. Species visiting trays in both yard designs had lower GUDs in mesic yards. Differences in resource abundance (i.e., alternative resources more abundant and of higher quality in xeric yards) contributed to our results, while predation costs associated with foraging did not. By enhancing the GUD, a common method for assessing the costs associated with foraging, our statistical model provided insights into how individual species and bird densities influenced the GUD. These differences we found in foraging behavior were indicative of differences in habitat quality, and thus our study lends additional support for native landscapes to help reverse the loss of urban bird diversity.     

Qualitative geographical variation in interspecific interactions

We explore geographical variation in the density relationship between potential competitor forest bird groups, resident Parus spp. and migrant Fringilla spp., across Europe using published bird census results. In addition, we summarized results from three experimental studies from northern Europe on their density associations. Based on anticipated changes in the relative intensity of positive and competitive interactions we predicted a unimodal density association between Parus and Fringilla : at low and intermediate densities the two groups are positively associated (positive interaction), whereas high densities promote interspecific competition. In central Europe where densities are high, densities were unimodally related to each other. In northern and southern Europe linear and positive associations appeared. Experimental studies provided consistent support for positive interspecific interactions in the north. The results suggest that species interactions may indeed vary in relation to the density of potential competitor and switch from positive to negative along environmental gradients.     

Opportunity costs influence food selection and giving‐up density of dabbling ducks

The interaction of animals with their food can yield insights into habitat characteristics, such as perceived predation risk and relative quality. We deployed experimental foraging patches in wetlands used by migrating dabbling ducks Anas spp. in the central Illinois River Valley to estimate variation in seed removal and giving‐up density (GUD; i.e. density of food remaining in patches following abandonment) with respect to seed density, seed size, seed depth in the substrate, substrate firmness, perceived predation risk, and an energetic profitability threshold (i.e. critical food density). Seed depth and the density of naturally‐occurring seeds outside of experimental plots affected seed removal and GUD in experimental patches more than perceived predation risk, seed density, seed size or substrate firmness. The greatest seed removal and lowest GUDs in experimental patches occurred when food resources in alternative foraging locations outside of plots (i.e. opportunity costs) appeared to be near or below a critical food density (i.e. 119–181 kg ha^–1). Giving‐up densities varied substantially from a critical food density across a range of food densities in alternative foraging locations suggesting that fixed GUDs should not be used as surrogates for critical food densities in energetic carrying capacity models. Foraging and resting rates in and near experimental foraging patches did not reflect patterns of seed removal and were poor predictors of GUD and foraging habitat quality. Our results demonstrated the usefulness of GUDs as indicators of habitat quality for subsurface, benthic foragers relative to other available foraging patches and suggested that food may be limited for dabbling ducks during spring migration in some years in the midwestern USA.     

Ontogenetic shifts in the functional response and interference interactions of Rhyacophila dorsalis larvae (Trichoptera)

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Therefore, it is important to establish if the functional response and interference interactions differ between life‐stages. These hypotheses were tested by (i) comparing the functional response of second, third, fourth and fifth larval instars of Rhyacophila dorsalis, using three stream tanks with one Rhyacophila larva per tank and one of 10 prey densities between 20 and 200 larvae of Chironomus sp.; (ii) using other experiments to assess interference within instars (two to five larvae of the same instar per tank), and between pairs of different instars (one, two or three larvae per instar; total predator densities of two, four or six larvae per tank). 2. The first hypothesis was supported. The number of prey eaten by each instar increased with prey density, the relationship being described by a type II model. The curvilinear response was stronger for fourth and fifth instars than for second and third instars. Mean handling time did not change significantly with prey density, and increased with decreasing instar number from 169 s for fifth instars to 200 s for second instars. Attack rate decreased progressively with decreasing instar number. Handling time varied considerably for each predator–prey encounter, but was normally distributed for each predator instar. Variations in attack rate and handling time were related to differences in activity between instars, fourth and fifth instars being more active and aggressive than second and third instars, and having a higher food intake. 3. The second hypothesis was partially supported. In the interference experiments between larvae of the same instar or different instars, mean handling time did not change significantly with increasing predator density, and attack rate did not change for second and third instars but decreased curvilinearly for fourth and fifth instars. Interference between some instars could not be studied because insufficient second instars were available at the same time as fourth and fifth instars, and most third instars were eaten by fourth and fifth instars in the experiments. Prey capture always decreased with decreasing attack rate. Therefore, interference reduced prey consumption in fourth and fifth instars, but not in second and third instars. The varying feeding responses of different instars should be taken into account when assessing their role in predator–prey relationships in the field.     

玉米螟赤眼蜂自身密度的干扰效应研究

玉米赤眼蜂自身密度干扰效应显著。在同一寄主（米蛾卵）密度条件下,干扰效应符合Hassell模型。在不同寄主密度条件下,赤眼蜂的寄生功能反应符合Beddington模型,自身密度对寄生率以及个体生殖力也存在干扰效应。