Foraging information affects the nature of competitive interactions

If animals can learn environmental cues, how might their use of information affect competitive interactions between species? I used shared‐preference isoleg theory to generate four different predictions regarding possible ways information could affect density‐dependent habitat selection. To test these predictions, I conducted field experiments on the foraging behavior of three coexisting species of hummingbirds. I studied black‐chinned (Archilochus alexandri), blue‐throated (Lampornis clemenciae), and magnificent (Eugenes fulgens) hummingbirds at the Southwestern Research Station in the Chiricahua Mts. of Southeastern Arizona, USA. Blue‐throated hummingbirds behaviorally dominate the other two species. I found that birds foraging with complete information (via learned color cues) avoid some of the negative effects from competition. Birds that foraged with complete information remained highly selective on rich feeders even with high competitor densities. When birds suffered an information deficit, however, the two subordinate species shifted their foraging preferences. The dominant species did not shift preference. Each subordinate's shift reflects its unique place in this competitively structured guild. With high competitor densities, black‐chinneds shifted from selective to opportunistic foraging when they also suffered from an information deficit. Thus, an information deficit caused black‐chinneds to cross over their first isoleg. Rather than reduce foraging efficiency, the magnificents shortened their foraging time. I use the magnificents’ response to competition and an information deficit to explore the possibility that this guild of hummingbirds is centrifugally organized. My results require us to consider the information‐gathering (e.g., learning) abilities of individual decision‐makers when we evaluate density‐dependent habitat selection. Differences in the level of usable information can alter the way coexisting species respond to competitive interactions that potentially structure communities.     

Influence of normal daytime fat deposition on laboratory measurements of torpor use in territorial versus nonterritorial hummingbirds

Fat deposition and torpor use in hummingbirds exhibiting distinct foraging styles should vary. We predicted that dominant territorial hummingbirds will use torpor less than subordinate nonterritorial species because unrestricted access to energy by territory owners allows for fat storage. Entry into torpor was monitored using open-flow respirometry on hummingbirds allowed to accumulate fat normally during the day. Fat accumulation was measured by solvent fat extraction. Territorial blue-throated hummingbirds (Lampornis clemenciae) had the highest fat accumulation and used torpor only 17% of the time. Fat storage by L. clemenciae averaged 26% of lean dry mass (LDM) in 1995 and 18% in 1996, similar to that measured for other nonmigratory birds. Fat storage by magnificent hummingbirds (Eugenes fulgens; trapliner) and black-chinned hummingbirds (Archilochus alexandri; nectar robber) averaged 19% and 16% of LDM, respectively, and they used torpor frequently (64% and 92% of the time, respectively). All species initiated torpor if total body fat dropped below 10% of LDM, indicating the existence of a torpor threshold. The ability of L. clemenciae to store enough fat to support nighttime metabolism is likely an important benefit of territoriality. Likewise, frequent torpor use by subordinates suggests that natural restrictions to energy intake can impact their energy budget, necessitating energy conservation by use of torpor.     

Discrimination of colors by the black-chinned hummingbird,Archilochus alexandri

Summary The color vision of a population of black-chinned hummingbirds was studied by behavioral methods. Birds were attracted to feeders equipped with tungsten lamps and interference filters. Results are based on counts of approximately 5700 visits by 92 ± 5 birds. Population size was estimated by mistnetting and marking 29 hummingbirds, 22 of which could be recognized individually during the course of the experiments.Following experience with red (620 nm) at all feeders, the birds showed a modest tendency to visit red (620 nm, 650 nm) and blue (490 nm) rather than intermediate greens and yellows. When sugar was presented at only one wavelength, however, choices became much sharper.When positions of the feeders were randomized, trained birds selected feeders on the basis of hue. Brightness was not used as a significant cue. This finding thus provides a more rigorous demonstration of color vision in hummingbirds than has heretofore been available.Either position or color could be learned in several hours (6–22 visits). Red (620 nm) and green (546 nm) were learned at the same rate. Two different (and opposing) color associations could be learned simultaneously at sites approximately 30 m apart.Discrimination of hue was measured following training to each of four wavelengths: 620, 590, 546, and 480 nm. Light from interference filters with transmission maxima at 546 and 550 nm were differentiated by the birds to a statistically significant extent. 546 and 590 nm appear to lie near the boundaries of hues; a boundary near 540 nm is found in pigeon but not human color vision.These experiments were supported by NIH grant EY00222. We are grateful to Nate Mandell for construction of the feeder-stimulators, to Vince Roth and the staff of the Southwestern Research Station for accommodating our special needs during the course of this work, and to the Bird Banding Laboratory of the Fish and Wildlife Service and the Arizona Fish and Game Department for granting the necessary permits to net and mark hummingbirds. We also thank Reed Pike of the Department of Structural Biology, Stanford University for unfailing help in preparing the illustrations.     

A wavelength discrimination function for the hummingbirdArchilochus alexandri

Summary Free-flying black-chinned hummingbirds (Archilochus alexandri) at a site in southeastern Arizona were attracted to artificial feeders displaying narrow spectral bands of light (7 nm half band width). The birds were taught to discriminate between pairs of wavelengths of approximately equal brightness but with a spectral separation of 10 nm. After training, performance of the birds was not significantly changed by alterations in the relative intensities of the two lights. Moreover, when the spectral composition of the test and training lights was made identical, the birds did not learn to make a discrimination on the basis of intensity differences of 0.5 or 1 log unit. In the learned foraging behavior of these hummingbirds, the salience of brightness is therefore inconsequential relative to hue.Discrimination scores for a constant 10 nm separation of test and training wavelengths were determined between 410 and 650 nm. This measure of the spectral dependence of wavelength discrimination shows a deterioration of performance at the red end of the spectrum but not in the blue and violet. Moreover, the minima at 585 and 555 nm indicate more structure than is present in this region of the spectrum in the human hue discrimination curve, and are similar but not identical to data on pigeon. These results are consistent with a growing body of evidence suggesting that the color space of birds may be more than three dimensional.This work was supported by NIH grants EY03266 and EY00222. We are indebted to Sally and Walter Spofford, who generously allowed us to work at their home, Aguila-Rancho, during May and June of 1980, and without whose kind help these experiments could not have been performed.     

Irrational choices in hummingbird foraging behaviour

It is conventionally assumed that, when animals evaluate alternative options, the value assigned to an option is absolute and independent of the other options available. It follows that animal choices should exhibit the rational property of regularity whereby the proportion of choices for an option cannot be increased by the addition of further options to the choice set. However, violations of regularity occur in human decision making, suggesting that humans may use comparative evaluation mechanisms whereby the value of an option is computed relative to the other options available. For example, in the asymmetrically dominated decoy effect the preference for a target option over a competitor is altered by the addition of a decoy option that is inferior to the target and competitor on one attribute, but lies between them on a second. We tested whether foraging wild rufous hummingbirds, Selasphorus rufus, would demonstrate violations of regularity in response to an asymmetrically dominated decoy. Sixteen birds chose between three artificial flower types (Target: 15 μl, 40% sucrose; Competitor: 45 μl, 30%; Decoy: 10 μl, 35%) in Binary (Target versus Competitor) and Trinary (Target versus Competitor versus Decoy) treatments. We predicted higher preference for the Target in the Trinary treatment. The birds ranked the three options in the same order in the Binary and Trinary treatments (Competitor>Target>Decoy). Seven birds showed violations of regularity, six increasing their absolute preference for the Competitor in the Trinary treatment. Overall, relative preference for the Competitor over the Target was higher in the Trinary than in the Binary treatment. These changes in preference are incompatible with an absolute evaluation mechanism.     

Investigations of density interactions among breeding birds in ponderosa pine forests: correlative and experimental evidence

Summary We assessed the importance of interspecific competition among insectivorous birds breeding in northcentral Arizona's ponderosa pine forests. We examined density interactions among species using two analytic approaches; correlative and experimental. The correlative approach examined patterns of change in breeding densities over four years at the community level and within two foraging guilds; picker-gleaners and aerial feeders. The relationships between morphological and behavioral similarity with pairwise density interactions were also assessed. Our experimental approach involved placement of nest boxes on two treatment plots to increase breeding densities of secondary cavity nesting birds that were in foraging guilds with open nesting insectivores.We found little evidence of interspecific competition. Patterns of density fluctuations indicated large positive covariances among species at both the community level and within guilds. Pairwise density interactions were independent of morphological or behavioral similarity. Nest boxes significantly increased breeding densities of the secondary cavity nesters. However, these increases did not induce reprocal density changes in the open nesting species. Interspecific competition for food during the breeding season appears to be unimportant in ponderosa pine bird communities.     

The foraging behavior of granivorous rodents and short-term apparent competition among seeds

The foraging behavior of a predator species is thought to bethe cause of short-term apparent competition among those preyspecies that share the predator. Short-term apparent competitionis the negative indirect effect that one prey species has onanother prey species via its effects on predator foraging behavior.In theory, the density-dependent foraging behavior of granivorousrodents and their preference for certain seeds are capable of inducing short-term apparent competition among seed species.In this study, I examined the foraging behavior of two heteromyidrodent species (family Heteromyidae), Merriam's kangaroo rats(Dipodomys merriami) and little pocket mice (Perognathus longimembris).In one experiment I tested the preferences of both rodent speciesfor the seeds of eight plant species. Both rodent species exhibiteddistinct but variable preferences for some seeds and avoidanceof others. However, the differences in preference appearedto have only an occasional effect on the strength of the short-term apparent competition detected in a field experiment. In anotherexperiment, I found that captive individuals of both rodentspecies had approximately equal foraging effort (i.e., timespent foraging) in patches that contained a highly preferredseed type (Oryzopsis hymenoides) regardless of seed density and the presence of a less preferred seed type (Astragalus cicer)in the patches. The rodents also harvested a large proportionof O. hymenoides seeds regardless of initial seed density;this precluded a negative indirect effect of A. cicer on O.hymenoides. But there was a negative indirect effect of O.hymenoides on A. cicer caused by rodents having a lower foragingeffort in patches that only contained A. cicer seeds than inpatches that contained A. cicer and O. hymenoides seeds. Theindirect interaction between O. hymenoides and A. cicer thusrepresented a case of short-term apparent competition thatwas non-reciprocal. Most importantly, it was caused by theforaging behavior of the rodents.     

Ultrasonic singing by the blue-throated hummingbird: a comparison between production and perception

Blue-throated hummingbirds produce elaborate songs extending into the ultrasonic frequency range, up to 30 kHz. Ultrasonic song elements include harmonics and extensions of audible notes, non-harmonic components of audible syllables, and sounds produced at frequencies above 20 kHz without corresponding hearing range sound. To determine whether ultrasonic song elements function in intraspecific communication, we tested the hearing range of male and female blue-throated hummingbirds. We measured auditory thresholds for tone pips ranging from 1 kHz to 50 kHz using auditory brainstem responses. Neither male nor female blue-throated hummingbirds appear to be able to hear above 7 kHz. No auditory brainstem responses could be detected between 8 and 50 kHz at 90 dB. This high-frequency cutoff is well within the range reported for other species of birds. These results suggest that high-frequency song elements are not used in intraspecific communication. We propose that the restricted hummingbird hearing range may exemplify a phylogenetic constraint.     

Experimental evidence for the use of density based interspecific social information in forest birds

Reproductive success and habitat preference are generally assumed to be negatively associated with densities of con- and heterospecific competitors. However, recent theoretical studies have suggested that in some cases habitat preference may have a nonlinear unimodal function in relation to con- or heterospecific competitor densities – intermediate densities being preferred. Such a pattern is expected if con- or heterospecific densities are used as a proximate cue in habitat selection, which may produce benefits by reducing searching costs and providing information about current habitat quality and costs of competition. At low density the use of such cues, and hence habitat selection, are hampered, whereas at high density costs of competition exceed the benefits of using cues, leading to avoidance. Here, we tested this hypothesis by examining whether arboreal migratory birds use the density of resident titmice ( Parus spp.) in habitat selection decisions. Many migrants and titmice species share similar resource needs making titmice density a reliable source of information for migrants. At the scale of habitat patches, we experimentally created a range of titmice densities from low to very high and subsequently measured the density response of migrants. In contrast to the unimodal habitat preference hypothesis, the average species number and total density of migratory birds were positively and linearly correlated with manipulated titmice density. Thus, migrants probably use titmice density as a relative indicator of habitat quality (abundance or quality of food) because foliage gleaners that share similar food resource with titmice, but not ground foragers, showed a positive association with manipulated titmice density. These results emphasize the positive effect of interspecific social information on habitat choice decisions and diversity of migratory bird community.     

What do foraging hummingbirds maximize?

Summary Hainsworth and Wolf (1976) reported that under certain conditions hummingbirds made food choices which did not maximize their net rate of energy intake while foraging. They concluded that the birds were not foraging optimally. We show here that their birds probably maximized a different utility function, the net energy per unit volume consumed (NEVC), which appears to be an optimal choice on a time scale longer than that of a foraging bout. Our own experiments with Archilochus colubris support the conclusion that hummingbirds make foraging decisions that maximize NEVC. A simulation model shows that, in nature, NEVC maximization would require fewer foraging trips and visits to fewer flowers per day to balance daily energy budgets. For territorial birds this can lead to smaller territory sizes and reduced costs of territorial defense. Plants that evolutionarily increase corolla length to enhance pollinator specificity need only increase nectar concentration slightly to maintain the same net energy per unit volume consumed (NEVC) by a given hummingbird pollinator.     

Consequences of differences in body mass,wing length and leg morphology for nectar-feeding birds

Nectar-feeding birds are prominent in many parts of the world, and vary with respect to body size. Despite the availability of considerable morphometric data, few concerted efforts have been made to assess the influence of attributes such as mass, wing length and leg morphology upon the speed, acceleration, mode and energetic cost of movement by birds between flowers when foraging for nectar. This review attempts to consolidate and interpret available data and highlight areas where further investigations appear warranted. Australian honeyeaters are generally larger, and American hummingbirds smaller, than Hawaiian honeycreepers and sunbirds of Africa or Asia. Sunbirds, honeyeaters and honeycreepers generally perch while extracting nectar from flowers. Hummingbirds usually hover, apparently because suitable perches close to flowers are lacking, and not because hovering increases the speed at which flowers can be visited. Honeyeaters move from one flower to another at speeds that are at least as great as those for hummingbirds. Most passerine nectarivores need to ingest more nectar per day than hummingbirds in order to maintain energy balance, some species devoting more than 60% of the day to foraging. The major consequence of reduced foraging activity by hummingbirds, which spend only 5–30% of the day in this manner, appears to be male emancipation from nest construction and care of offspring. Large nectarivores have a greater capacity to store surplus food and to fast than smaller birds, and so can take advantage of short-lived peaks in nectar abundance. Nectarivores such as honeyeaters should therefore be favoured by the rapid diurnal changes in nectar availability which are characteristic of many Australian and African habitats. Body mass also determines the likely access to rich sources of nectar through size-related interspecific dominance hierarchies. In all families, larger species tend to monopolize the most rewarding nectar supplies, forcing smaller subordinate species to use poorer, more scattered sources. Within particular species, males usually have longer wings and greater masses than females. These variations imply that the two sexes differ with regard to their foraging ecology, although few supporting data are currently available.     

Pairing automated mark–recapture and social network models to explore the effects of landscape configuration on hummingbird foraging patterns

Landscape changes can alter pollinator movement and foraging patterns which can in turn influence the demographic processes of plant populations. We leveraged social network models and four fixed arrays of five hummingbird feeders equipped with radio frequency identification (RFID) data loggers to study rufous hummingbird (Selasphorus rufus) foraging patterns in a heterogeneous landscape. Using a space-for-time approach, we asked whether forest encroachment on alpine meadows could restrict hummingbird foraging movements and impede resource discovery. We fit social network models to data on 2221 movements between feeders made by 29 hummingbirds. Movements were made primarily by females, likely due to male territoriality and early migration dates. Distance was the driving factor in determining the rate of movements among feeders. The posterior mean effects of forest landscape variables (local canopy cover and intervening forest cover) were negative, but with considerable uncertainty. Finally, we found strong reciprocity in hummingbird movements, indicative of frequent out and back movements between resources. Together, these findings suggest that reciprocal movements by female hummingbirds could help maintain bidirectional gene flow among nearby subpopulations of ornithophilous plants; however, if the distance among meadows increases with further forest encroachment, this may limit foraging among progressively isolated meadows.     

Information for patch assessment: a field investigation with black-chinned hummingbirds

I examined the kinds of information black-chinned hummingbirds(Archilochus alexandri) used when exploiting artificial resourcepatches in the field. I determined whether birds (1) only usedcurrent patch-sample information or (2) combined current patch-samplewith prior information (the distribution of resources amongpatches) to estimate patch quality, and (3) whether the kindsof information used depended on environmental variability. Iestablished two environments that differed in amount of variationin patch subtypes and determined the use of information by assessingthe correlation between the number of rewards obtained and thenumber of consecutive unrewarded patch probes just before patchdeparture. In the low-variance environment, most birds appearedto combine prior information with patch-sample information.Individuals using prior information foraged more efficientlythan those that did not. In the high-variance environment, somebirds appeared to combine patch-sample information with priorinformation, whereas others apparently relied solely on patch-sampleinformation to estimate patch quality, with no difference inforaging efficiency. In both environments, prior informationwas used after approximately 25 patches were exploited.     

Variation in pollinator preference between two Ipomopsis contact sites that differ in hybridization rate

Pollinator-mediated reproductive isolation is often a principal factor in determining the rate of hybridization between plant species. Pollinator preference and constancy can reduce interspecific pollen transfer between otherwise interfertile, coflowering species. The importance of this ethological isolation can be assessed by comparing the strength of preference and constancy of pollinators in contact sites that differ in the frequency of hybrid individuals. We observed visitation by hummingbirds and hawkmoths in natural single-species patches and artificial mixed-species arrays in two Ipomopsis aggregata/I. tenuituba contact sites-one with few hybrids, and one in which hybrids are abundant. Pollinator preference and constancy were stronger at the low-frequency hybrid site, especially for hawkmoths (Hyles lineata). Hawkmoths at the low-frequency hybrid site showed significant preference and constancy for I. tenuituba, while at the high-frequency site hawkmoths visited both species equally. One hypothesis that might explain these differences in hawkmoth foraging is that warmer nights at the low-frequency hybrid site allow for nocturnal foraging where the light-colored corollas of I. tenuituba have a visibility advantage. These differences in hawkmoth behavior might in turn affect hummingbirds differently at the two sites, through changes in nectar resources, leading to greater pollinator-mediated isolation at the low-frequency hybrid site. Our results suggest that differences in pollinator behaviors between sites can have both direct and indirect effects on hybridization rates between plant species.     

Disentangling olfactory and visual information used by field foraging birds

Foraging strategies of birds can influence trophic plant–insect networks with impacts on primary plant production. Recent experiments show that some forest insectivorous birds can use herbivore‐induced plant volatiles (HIPVs) to locate herbivore‐infested trees, but it is unclear how birds combine or prioritize visual and olfactory information when making foraging decisions. Here, we investigated attraction of ground‐foraging birds to HIPVs and visible prey in short vegetation on farmland in a series of foraging choice experiments. Birds showed an initial preference for HIPVs when visual information was the same for all choice options (i.e., one experimental setup had all options with visible prey, another setup with hidden prey). However, if the alternatives within an experimental setup included visible prey (without HIPV) in competition with HIPV‐only, then birds preferred the visual option over HIPVs. Our results show that olfactory cues can play an important role in birds’ foraging choices when visual information contains little variation; however, visual cues are preferred when variation is present. This suggests certain aspects of bird foraging decisions in agricultural habitats are mediated by olfactory interaction mechanisms between birds and plants. We also found that birds from variety of dietary food guilds were attracted to HIPVs; hence, the ability of birds to use plant cues is probably more general than previously thought, and may influence the biological pest control potential of birds on farmland.     

Patterns of resource use in an assemblage of birds in the canopy of a temperate alluvial forest

In our study, we assessed patterns of resource use in an assemblage of birds by observing their foraging behaviour from a crane in the canopy of a temperate alluvial forest. We selected 12 bird species and addressed seasonal changes in feeding activity during a 2-month period in spring focussing on average staying time and utilisation of crown strata in two tree species, the common oak (Quercus robur) and the sycamore maple (Acer pseudoplatanus). We further examined ecological characteristics of the trees (i. e., crown density) that are likely to influence resource use in birds. The selected birds differed in their preference for the tree species. Most birds preferred common oaks. This preference was probably associated with higher food abundance related to substrate characteristics (i.e., roughness of bark) which offer more microhabitats for arthropods and thus permit higher densities of potential prey. Some bird species switched feeding preferences within the study period from sycamore maples to common oaks in association with tree phenology. We found two main foraging techniques. All birds searched for prey at short distance (≤50 cm) and gleaned food from substrate except the Pied Flycatcher that foraged by hovering and searched over longer distances (>50 cm). Overall, we demonstrate in our study that canopy access with mobile crane systems provides excellent opportunities to observe canopy birds and enables detailed analysis of their foraging behaviour. The main result of our study reveals fine-grained resource partitioning of birds within the canopy as an important factor structuring assemblages, with species-specific and in part also seasonal differences in stratification and substrate use.     

Oriole pollination ofErythrina breviflora (Leguminosae): Evidence for a polytypic view of ornithophily

Erythrina breviflora is visited by large numbers of passerine birds of which orioles (Icterus: Icteridae) are the primary pollinators. The flowers produce large quantities of nectar but they are rarely visited by hummingbirds. Inflorescence and floral morphology, and low levels of sucrose in the nectar probably explain the rarity of foraging hummingbirds. A comparison of Old WorldErythrina and their pollinators with New World species pollinated by orioles and hummingbirds suggests that parallel evolution has occurred. When the comparison is expanded to include other species pollinated by orioles, it is clear that various New WorldIcteridae, Thraupidae, etc. are ecological equivalents of Old WorldOriolidae, Pycnonotidae, Sturnidae, etc. and that flowers pollinated by these birds have similar characteristics.     

Effects of competitor density and physical habitat structure on the competitive intensity of territorial white spotted charr Salvelinus leucomaenis

This study compared the effects of interference competition in habitats of different complexity and in different densities. The influence of fish density and habitat structure was examined in manipulative experiments using young-of-the-year white spotted charr Salvelinus leucomaenis as a model species. The difference of specific growth rate ( G ) range, an index of interference competitive intensity, was significantly smaller in the structurally complex treatments than structurally simple treatments, while there were no significant difference between high-density and low-density treatments. Thus, physical habitat structure was more effective for mitigating interference competition than manipulating competitor density. Although interference competition was not affected by competitor density, mean G were suppressed in the high-density treatments. This implied that exploitative competition may cause the decrease of G rather than interference competition does in the high-density treatments. Mean G were also suppressed in the structurally complex treatments. Chaotic flow pattern created by physical habitat structures may decrease G by reducing foraging success of experimental fish in the complex treatments.     

Context-dependent foraging decisions in rufous hummingbirds

A core assumption implicit in economic models of animal choice is that subjects assign absolute utilities to options that are independent of the type and number of alternatives available. Humans sometimes appear to violate this assumption and employ relative, as opposed to absolute, currencies when making choices. Recent evidence suggests that animals too might sometimes employ relative choice mechanisms. We tested this idea by measuring the foraging preferences of rufous hummingbirds (Selasphorus rufus) faced with choices analogous to those in which human use of relative currencies is evident. The birds experienced three treatments: a binary choice between two artificial flower types designated concentration (20 microl, 40% sucrose solution) and volume (40 microl, 20%), and two trinary treatments in which a third decoy option (either concentration decoy: 10 microl, 30% or volume decoy: 30 microl, 10%) was added to the set. The birds' preferences differed significantly across the three treatments. In the trinary treatments, the effect of the decoy options was to increase the preference for the option that dominated the decoy. These results are similar to those reported in the human choice literature, and are compatible with the hummingbirds using a relative evaluation mechanism in decision making.     

Spatial clumping of food and social dominance affect interference competition among ruddy turnstones

In studying the success of foraging animals, studies of interferencecompetition have put emphasis on effects of competitor density,whereas studies of resource defense have focused on the effectsof the spatial distribution of food within patches. Very fewstudies have looked at both factors simultaneously, that is,determined whether the effects of competitor density on foragingsuccess depend on the spatial distribution of food. We studiedthe behavior and the foraging success of ruddy turnstones (Arenariainterpres) using an experiment in which we varied both the presenceof a competitor and the food distribution. Because turnstonesmay differ strongly in their relative dominance status, we alsoexperimentally varied the foragers' relative dominance status.We found that the presence of a competitor only reduced theforaging success of subordinate birds foraging at the clumpedfood distribution. At this condition, dominant and subordinatebirds differed markedly in their foraging success. Contraryto our expectations, we did not observe more agonistic behaviorat the clumped food distribution. This indicates that the amountof agonistic behavior observed may be a bad indicator of interferenceeffects. These findings have specific implications for modelsof interference competition. Most notably they show that theeffects of competitor density on agonistic behavior and foragingsuccess may well depend on the spatial distribution of foodand the foragers' relative dominance status. Additionally, ourresults suggest that social dominance will not be fully understoodwithout considering long-term processes such as the formationand maintenance of social dominance hierarchies.