The intensity of interference varies with food density: support for behaviour-based models of interference

Dolman (1995) measured the intake rates of snow buntings feeding on seed patches at difference bird densities, for each of two different seed densities. Interference occurred in the low food density treatment, with intake rate declining at high bird densities, but did not occur in the high food density treatment. Dolman states that existing models of the interference process assume that the intensity of interference is independent of food density. While this statement is true for the model of Hassell and Varley (1969), we show that behaviour-based models of interference do not assume that interference is independent of food density. We examine two simple analytic behaviour-based models of interference and show that, in agreement with Dolman's observations, intensity of interference is predicted to decrease with increasing food density. Dolman plotted log (intake rate) against log (bird density). Rather than obtaining a linear relationship predicted by the Hassell and Varley model, the results from the low seed density treatment indicated a curvilinear relationship consistent with those produced by our behaviour-based models. These results provide support for the use of behaviour-based models of interference over the model of Hassell and Varley.     

A shift from exploitation to interference competition with increasing density affects population and community dynamics

Intraspecific competition influences population and community dynamics and occurs via two mechanisms. Exploitative competition is an indirect effect that occurs through use of a shared resource and depends on resource availability. Interference competition occurs by obstructing access to a resource and may not depend on resource availability. Our study tested whether the strength of interference competition changes with protozoa population density. We grew experimental microcosms of protozoa and bacteria under different combinations of protozoan density and basal resource availability. We then solved a dynamic predator–prey model for parameters of the functional response using population growth rates measured in our experiment. As population density increased, competition shifted from exploitation to interference, and competition was less dependent on resource levels. Surprisingly, the effect of resources was weakest when competition was the most intense. We found that at low population densities, competition was largely exploitative and resource availability had a large effect on population growth rates, but the effect of resources was much weaker at high densities. This shift in competitive mechanism could have implications for interspecific competition, trophic interactions, community diversity, and natural selection. We also tested whether this shift in the mechanism of competition with protozoa density affected the structure of the bacterial prey community. We found that both resources and protozoa density affected the structure of the bacterial prey community, suggesting that competitive mechanism may also affect trophic interactions.     

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.     

Regulatory Mechanisms of Reproductive Effort in Plants II. Plasticity in Reproductive Energy Allocation and Propagule Output of Glycine max Merr. (Leguminosae) Cultivated at Varying Densities and Nitrogen Levels

Abstract Plasticity in growth, reproductive energy allocation (RA), and reproductive output were studied in Glycine max Merr. Cv. Enrei (Leguminosae) grown under varying densities and soil nitrogen levels.
Marked plastic responses were detected in individual biomass, the patterns of resource allocation to total reproductive structures (RA) and also to propagules, reproductive outputs, and propagule weight under changing densities and soil nitrogen levels. Plants cultivated at higher densities exhibited proportionately lower individual biomass, lower RA, lower seed output, and smaller seed size in response to increasing density and decreasing soil nitrogen levels, although some deviations were observed, especially in the highest density plot with no fertilization. Differences due to different N-levels were not as great as those to changing density, which may in part be due to the fact that soybean has nitrogen-fixing bacteria in root tubercles, just as in any other Leguminosae. Fecundity was also maintained at the similar high rates of 80–97% in all plots examined, although slight but steady decreases were noted with increasing density. This resemblance in fecundity may be due to its strong inbreeding system.
Another important finding was that seed production under limited resource availability, notably lack of ample solar radiation due to strong interference at higher density plots, is exceedingly costly. This was most clearly exhibited by a sharp increase in relative energy partitioning to a single propagule in response to the increased density, the relative energy cost to a single propagule (RA) increasing from one to seven-fold. The results obtained in this study coincide well with the findings made in other plants, e.g., Helianthus annuus, Oryza sativa , and Coix ma-yuen , with the same experimental designs.     

Is the pollen‐limited mistletoe Peraxilla tetrapetala (Loranthaceae) also seed limited?

Abstract Seed production in the endemic New Zealand mistletoe Peraxilla tetrapetala (Loranthaceae) has been shown to be consistently pollen limited, but to date there has been no test of whether the species is also seed limited. If it is not seed limited, then pollen limitation may have no effect on population size. We tested for seed limitation by sowing seeds onto host trees at high and low densities at two sites, and following survival for up to 69 months. Our rationale was that unless there is strong density‐dependent seedling mortality sufficient to negate increases in seed supply, the species would be seed limited. We simultaneously measured pollen limitation at both sites using pollen augmentation. Peraxilla tetrapetala was strongly pollen limited at both sites over 7 years at Ohau (mean Pollen Limitation Index 0.62), and 12 years at Craigieburn (mean PLI 0.44), before and during the seed‐limitation study. There was no significant overall negative effect of density on survival in sown P. tetrapetala seeds over 42–69 months. There was a significant positive effect: seeds were more likely to adhere to branches when sown at high densities. This initial advantage to high‐density seeds was gradually eroded by slightly (but non‐significantly) lower survival rates of adhered seeds at high density. By the end of the study there was no significant difference in overall survival in high and low density plantings. This means that P. tetrapetala was apparently both pollen limited and seed limited at both our sites. Hence, reduced densities of native bird pollinators caused by introduced mammalian carnivores are likely to reduce the density of adult mistletoes in the next generation. However, the generality of this result may be affected by the fact that mistletoes do not have a dense ‘seedling shadow’ under the parent mistletoe.     

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.     

Linking optimal foraging behavior to bird community structure in an urban-desert landscape: field experiments with artificial food patches

Urban bird communities exhibit high population densities and low species diversity, yet mechanisms behind these patterns remain largely untested. We present results from experimental studies of behavioral mechanisms underlying these patterns and provide a test of foraging theory applied to urban bird communities. We measured foraging decisions at artificial food patches to assess how urban habitats differ from wildlands in predation risk, missed-opportunity cost, competition, and metabolic cost. By manipulating seed trays, we compared leftover seed (giving-up density) in urban and desert habitats in Arizona. Deserts exhibited higher predation risk than urban habitats. Only desert birds quit patches earlier when increasing the missed-opportunity cost. House finches and house sparrows coexist by trading off travel cost against foraging efficiency. In exclusion experiments, urban doves were more efficient foragers than passerines. Providing water decreased digestive costs only in the desert. At the population level, reduced predation and higher resource abundance drive the increased densities in cities. At the community level, the decline in diversity may involve exclusion of native species by highly efficient urban specialists. Competitive interactions play significant roles in structuring urban bird communities. Our results indicate the importance and potential of mechanistic approaches for future urban bird community studies.     

The control of rank-abundance distributions by a competitive despotic species

Accounting for differences in abundances among species remains a high priority for community ecology. While there has been more than 80 years of work on trying to explain the characteristic S shape of rank-abundance distributions (RADs), there has been recent conjecture that the form may not depend on ecological processes per se but may be a general phenomenon arising in many unrelated disciplines. We show that the RAD shape can be influenced by an ecological process, namely, interference competition. The noisy miner (Manorina melanocephala) is a hyperaggressive, ‘despotic’ bird that occurs over much of eastern Australia (>10⁶ km²). We compiled data for bird communities from 350 locations within its range, which were collected using standard avian survey methods. We used hierarchical Bayesian models to show that the RAD shape was much altered when the abundance of the strong interactor exceeded a threshold density; RADs consistently were steeper when the density of the noisy miner ≥2.5 birds ha^?1. The structure of bird communities at sites where the noisy miner exceeded this density was very different from that at sites where the densities fell below the threshold: species richness and Shannon diversity were much reduced, but mean abundances and mean avian biomass per site did not differ substantially.     

Native Birds and Alien Insects: Spatial Density Dependence in Songbird Predation of Invading Oak Gallwasps

Revealing the interactions between alien species and native communities is central to understanding the ecological consequences of range expansion. Much has been learned through study of the communities developing around invading herbivorous insects. Much less, however, is known about the significance of such aliens for native vertebrate predators for which invaders may represent a novel food source. We quantified spatial patterns in native bird predation of invading gall-inducing Andricus wasps associated with introduced Turkey oak (Quercus cerris) at eight sites across the UK. These gallwasps are available at high density before the emergence of caterpillars that are the principle spring food of native insectivorous birds. Native birds showed positive spatial density dependence in gall attack rates at two sites in southern England, foraging most extensively on trees with highest gall densities. In a subsequent study at one of these sites, positive spatial density dependence persisted through four of five sequential week-long periods of data collection. Both patterns imply that invading galls are a significant resource for at least some native bird populations. Density dependence was strongest in southern UK bird populations that have had longest exposure to the invading gallwasps. We hypothesise that this pattern results from the time taken for native bird populations to learn how to exploit this novel resource.     

Regulatory Mechanisms of Reproductive Effort in Plants 1. Plasticity in Reproductive Energy Allocation and Propagule Output of Helianthus annuus L. (Compositae) Cultivated at Varying Densities and Nitrogen Levels

Abstract Plasticity in growth, reproductive energy allocation (RA), and reproductive output were investigated in Helianthus annuus L. cultivar. Russia (Compositae) grown under varying densities and soil nitrogen levels. The role and behaviour of pollinators in seed production was also examined.
Exceedingly marked plastic responses were detected in individual biomass, the patterns of resource allocation to total reproductive structures (RA) and also to propagules, fecundity, reproductive outputs, and propagule size and weight under changing densities and soil nitrogen levels. Plants cultivated at higher densities exhibited proportionately lower individual biomass, lower RA, lower fecundity, lower seed output, and smaller seed size in response to increasing density and decreasing soil nitrogen levels. However, differences due to different N-levels were not as great as those to changing density.
One of the most significant findings was that seed production under limited resource availability, i.e., lack of ample solar radiation and soil nutrients, due to strong interference at higher density plots, is exceedingly costlty , This was most clearly demonstrated by a very sharp increase in relative energy partitioning to a single propagule in response to the increased density and decreasing nitrogen levels, the relative energy cost to a single achcne (R_A) increasing from one to twenty-fourfold. Reproduction was also affected by pollinator-limitations, and seed size showed a marked position effect.     

Density-dependent habitat selection in migratory passerines during stopover: what causes the deviation from IFD?

We studied the distribution of migratory warblers (genus: Sylvia) in poor and high quality habitat patches at a stopover site in the northern Negev, Israel. The purpose of our study was to test predictions based on the ideal free distribution (IFD) model by using a natural ecosystem which has a high turnover of individuals moving between unfamiliar foraging patches. We trapped birds in two groves of Pistacia atlantica embedded within a coniferous forest. The fruit-density ratio between these groves was 45:1. We compared bird density, body condition and habitat matching (the ratio between bird density and resource density) at the two sites. To analyse the data we integrated two approaches to density-dependent habitat selection: the isodar method and the habitat matching rule. As predicted by the IFD model, we found that habitat suitability decreased with bird density with a high correlation between warbler densities in the two habitat patches. Contrary to IFD predictions, warbler density in the poor patch was higher than expected by the habitat-matching rule. This habitat under-matching, had a cost: in the rich habitat the average energy gain per individual bird was higher than in the poor habitat. Further analysis suggests that the apparent habitat under-matching is not due to interference or differences in warbler competitive abilities. Therefore, we suggest that this migratory bird community is not at equilibrium because the birds possess imperfect knowledge of resource distribution. We propose that this lack of knowledge leads to free, but not ideal distributions of migrant birds in unfamiliar stop over sites.     

Anthropogenically-Mediated Density Dependence in a Declining Farmland Bird

Land management intrinsically influences the distribution of animals and can consequently alter the potential for density-dependent processes to act within populations. For declining species, high densities of breeding territories are typically considered to represent productive populations. However, as density-dependent effects of food limitation or predator pressure may occur (especially when species are dependent upon separate nesting and foraging habitats), high territory density may limit per-capita productivity. Here, we use a declining but widespread European farmland bird, the yellowhammer Emberiza citrinella L., as a model system to test whether higher territory densities result in lower fledging success, parental provisioning rates or nestling growth rates compared to lower densities. Organic landscapes held higher territory densities, but nests on organic farms fledged fewer nestlings, translating to a 5 times higher rate of population shrinkage on organic farms compared to conventional. In addition, when parental provisioning behaviour was not restricted by predation risk (i.e., at times of low corvid activity), nestling provisioning rates were higher at lower territory densities, resulting in a much greater increase in nestling mass in low density areas, suggesting that food limitation occurred at high densities. These findings in turn suggest an ecological trap, whereby preferred nesting habitat does not provide sufficient food for rearing nestlings at high population density, creating a population sink. Habitat management for farmland birds should focus not simply on creating a high nesting density, but also on ensuring heterogeneous habitats to provide food resources in close proximity to nesting birds, even if this occurs through potentially restricting overall nest density but increasing population-level breeding success.     

Intraspecific and phylogenetic density-dependent seedling recruitment in a subtropical evergreen forest

Recent evidence suggests that plant performance can be influenced by the phylogenetic diversity of neighboring plants. However, no study to date has examined the effect of such phylogenetic density dependence on the transition from seed to seedling. Using 6 years of data on seedling recruitment and seed rain of 13 species from 130 stations (one 0.5 m² seed trap and three adjacent 1 m² seedling plots) in a subtropical evergreen forest, we asked: (1) Does negative density dependence act across seed to seedling stages? (2) Is there evidence for phylogenetic density dependence during the seed to seedling transition? (3) Does the strength of density dependence vary among years? Generalized linear mixed-effects models were used to model seed to seedling transition as a function of conspecific seed and seedling densities, heterospecific seed and seedling densities, and mean phylogenetic distance of heterospecific seeds and seedling. Conspecific seed density had a significant negative effect on seedling transition rates for 12 of 13 focal species. In contrast, conspecific seedling density had a positive effect for 7 species, suggesting species-specific habitat preferences. Few species were significantly affected by the density or phylogenetic relatedness of heterospecific seeds and seedlings. Only conspecific seed density effects varied among years for most focal species. Overall, our results reveal that conspecific seed and seedling densities play a more important role than the density or relatedness of heterospecific seeds and seedlings during the seed to seedling stage, suggesting that species-specific seed predators, along with habitat preferences, may contribute to diversity maintenance in this forest.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

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.     

Assessing the potential for Grey Squirrels Sciurus carolinensis to compete with birds at supplementary feeding stations

Supplementary feeding of birds, particularly in urban areas, is often associated with increased population size and fecundity. In the UK, the non‐native Grey Squirrel Sciurus carolinensis is common in rural and urban habitats. It exploits supplementary feeders and may induce interference competition by excluding birds, but empirical evidence of this is unavailable. Using controlled model presentation experiments, we demonstrate that Grey Squirrels could reduce bird use of supplementary feeders and induce interference competition. Total bird resource use was reduced by 98% and most species exhibited similar sensitivities. The likelihood and magnitude of interference competition will depend on how rapidly displaced birds find alternative food sources; it will be greatest where there are high Grey Squirrel densities and few supplementary feeders. Other studies suggest that supplementary feeding increases Grey Squirrel numbers, and the species is also predicted to expand its non‐native range across most of Europe. Our data indicate that Grey Squirrels may eventually alter the net effect of supplementary feeding on bird populations across the European continent; increased use of squirrel‐proof feeders may help to minimize such effects.     

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.     

Predictable changes in predation mortality as a consequence of changes in food availability and predation risk

Theory predicts that animals will have lower activity levels when either the risk of predation is high or the availability of resources in the environment is high. If encounter rates with predators are proportional to activity level, then we might expect predation mortality to be affected by resource availability and predator density independent of the number of effective predators. In a factorial experiment, we tested whether predation mortality of larval wood frogs, Rana sylvatica, caused by a single larval dragonfly, Anax junius, was affected by the presence of additional caged predators and elevated resource levels. Observations were consistent with predictions. The survival rate of the tadpoles increased when additional caged predators were present and when additional resources were provided. There was no significant interaction term between predator density and food concentration. Lower predation rates at higher predator density is a form of interference competition. Reduced activity of prey at higher predator density is a potential general mechanism for this widespread phenomenon. Higher predation rates at low food levels provides an indirect mechanism for density-dependent predation. When resources are depressed by elevated consumer densities, then the higher activity levels associated with low resource levels can lead to a positive association between consumer density and consumer mortality due to predation. These linkages between variation in behaviour and density-dependent processes argue that variation in behaviour may contribute to the dynamics of the populations. Because the capture rate of predators depends on the resources available to prey, the results also argue that models of food-web dynamics will have to incorporate adaptive variation in behaviour to make accurate predictions.     

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.     

Density- and frequency-dependent predation of artificial bird nests

Predators are often expected to vary their relative predation rates according to the frequency of prey types in the environment (frequency-dependent predation). The underlying cause for this must lie in some dependency of absolute predation rates on the density of prey types in the environment (density-dependent predation). However, frequency-dependent predation may either be caused by 'simple' density-dependent predation, in which the absolute predation rate on a given prey type depends purely on the density of that type, or by more complex responses in which absolute rates depend also on the density of other prey types. It is usually difficult to distinguish the underlying cause of frequency-dependent predation, because frequencies tend to change as densities change. Here, we describe the results of an experiment conducted to disentangle these phenomena under two prey richness (low and high) conditions. We used artificial bird nests (placed on shrubs and on saplings) baited with quail eggs placed in natural forests as models of natural bird nests. Our results indicate that both the absolute and relative predation rates on the prey types may vary in complex ways. Predation rates depend on a complex interaction between the prey's own density, other prey density and the diversity of prey in the environment. Neglecting to include, or consider, these complexities into analyses may lead to erroneous conclusions in studies of absolute or relative predation rates.