Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Asymmetric Competition Among Distant Taxa

For purposes of theoretical analysis, competition between distantlyrelated taxa is interpreted as asymmetric competition. Severalkinds of situations are examined: A predator competes with oneof its prey species for a second resource; competitors utilizesuccessive stages in the life cycles of prey; one predator facilitatesthe resource utilization of its competitor; competitors differin their vulnerability to predation. Methods of signed digraphs(loop analysis) and statistics over time are used to predictthe qualitative consequences of the different interaction patterns.     

Evolution of fighting behavior under asymmetric competition: an experimental test with juvenile salmonids

Large-dominant and small-subordinate species engaging in asymmetricinterference competition may optimize behavior under differenttrade-offs between the chance of winning and the cost of fighting.If fighting behavior is heritable and under selection, theorysuggests that large-dominant and small-subordinate species shouldevolve aggressive and passive fighting behaviors, respectively.To test this prediction, I manipulated the size and competitiveasymmetry of juveniles from sympatric populations of large-dominantcoho salmon (Oncorhynchus kisutch) and small-subordinate steelheadtrout (O. mykiss) and asked whether differences in fightingbehavior persisted independently of competitive ability. I observedfighting behavior during dyadic contests in two habitats, mutuallypreferred pools and energetically demanding riffles, under eachof three size treatments: natural size asymmetry, asymmetryremoved, and reversed size asymmetry. The results supportedthe prediction. Competitive ability depended primarily on size;large individuals of both species dominated smaller heterospecifics,and neither species dominated when size matched. Fighting behaviordepended primarily on species identity; coho salmon used a higherproportion of aggressive chases, whereas steelhead trout useda higher proportion of passive displays. Large individuals weremore likely to chase, and small individuals were more likelyto display. As evidence that asymmetric competition is associatedwith behavioral divergence, these results complement previouswork on morphological divergence under asymmetric competitionand provide a richer context for other features of the coho–steelheadsystem.     

OTHER PAPERS: INTERFERENCE AND RESOURCE COMPETITION IN TWO LAND SNAILS: ADULTS INHIBIT CONSPECIFIC JUVENILE GROWTH IN FIELD AND LABORATORY

Interference and resource competition by adults inhibited growthrates of conspecific juveniles of the land snail species Mesodonthyroidus and Neohelix albolabris in separate field and laboratoryexperiments, but not in laboratory experiments on Anguispiraalternata. In 1 m² field cages at near-natural densities underambient food and water conditions, juvenile M. thyroidus apparentlycompeted with adults for food or water or both resources, growingmore slowly when living with two conspecific adults, but beingunaffected by adult presence when food and water were augmented.Neohelix albolabris juveniles were similarly unaffected in fieldcages by presence of two adults when food and water were augmented.In contrast, interference, not resource competition, apparentlyexplained growth inhibition in laboratory cages at densitiesconsiderably greater than natural densities, with non-limitingfood and moisture; both M. thyroidus and N. albolabris juvenilesgrew more slowly as conspecific adult number increased fromzero to three. (Received 17 July 1995; accepted 11 November 1996)     

Two-species asymmetric competition: effects of age structure on intra- and interspecific interactions

1. The patterns of density-dependent resource competition and the mechanisms leading to competitive exclusion in an experimental two-species insect age-structured interaction were investigated. 2. The modes of competition (scramble or contest) and strength of competition (under- to overcompensatory) operating within and between the stages of the two species was found to be influenced by total competitor density, the age structure of the competitor community and whether competition is between stages of single or two species. 3. The effect of imposed resource limitation on survival was found to be asymmetric between stages and species. Environments supporting both dominant and subordinate competitors were found to increase survival of subordinate competitors at lower total competitor densities. Competitive environments during development within individual stage cohorts (i.e. small or large larvae), differed from the competitive environment in lumped age classes (i.e. development from egg-->pupae). 4. Competition within mixed-age, stage or species cohorts, when compared with uniform-aged or species cohorts, altered the position of a competitive environment on the scramble-contest spectrum. In some cases the competitive environment switched from undercompensatory contest to overcompensatory scramble competition. 5. Such switching modes of competition suggest that the relative importance of the mechanisms regulating single-species population dynamics (i.e. resource competition) may change when organisms are embedded within a wider community.     

Asymmetric interspecific competition drives shifts in signalling traits in fan-throated lizards

Interspecific competition can occur when species are unable to distinguish between conspecific and heterospecific mates or competitors when they occur in sympatry. Selection in response to interspecific competition can lead to shifts in signalling traits—a process called agonistic character displacement. In two fan-throated lizard species—Sitana laticeps and Sarada darwini—females are morphologically indistinguishable and male agonistic signalling behaviour is similar. Consequently, in areas where these species overlap, males engage in interspecific aggressive interactions. To test whether interspecific male aggression between Si. laticeps and Sa. darwini results in agonistic character displacement, we quantified species recognition and signalling behaviour using staged encounter assays with both conspecifics and heterospecifics across sympatric and allopatric populations of both species. We found an asymmetric pattern, wherein males of Si. laticeps but not Sa. darwini showed differences in competitor recognition and agonistic signalling traits (morphology and behaviour) in sympatry compared with allopatry. This asymmetric shift in traits is probably due to differences in competitive abilities between species and can minimize competitive interactions in zones of sympatry. Overall, our results support agonistic character displacement, and highlight the role of asymmetric interspecific competition in driving shifts in social signals.     

Predatory behavior and preference of a successful invader, the mud crab Dyspanopeus sayi (Panopeidae), on its bivalve prey

Predator-prey relationships between the panopeid crab, Dyspanopeus sayi, and the mytilid, Musculista senhousia, were investigated. Through laboratory experiments, prey-handling behavior, prey size selection, predator foraging behavior and preferences for two types of prey (M. senhousia and the Manila clam Ruditapes philippinarum) were assessed. Handling time differed significantly with respect to the three prey sizes offered (small: 15.0-20.0 mm shell length, SL; medium: 20.1-25.0 mm SL; and large: 25.1-30.0 mm SL); mud crabs were more efficient in predating medium-small than large prey. Although differences in prey profitability were not evident, D. sayi exhibited a marked reluctance to feed on larger-sized prey whilst smaller, more easily predated mussels were available. Size selection may be the result of a mechanical process in which encountered prey are attacked but rejected if they remain unbroken after a certain number of opening attempts. D. sayi exhibited inverse density-dependent foraging. A significant higher mortality of prey was evident at low prey density. Thus, at low predator density, the D. sayi-M. senhousia interaction was a destabilizing type II functional response. Interference responses affected the magnitude of predation intensity by D. sayi on M. senhousia, since as the density of foraging crabs increased, their foraging success fell. At high density (4 crabs tank⁻¹), crabs engaged in a high amount of agonistic activity when encountering a conspecific specimen, greatly diminished prey mortality. Finally, presenting two types of prey, Manila clam juveniles were poorly predated by mud crabs, which focused their predation mostly on M. senhousia. It is hypothesized that, when more accessible prey is available, mud crabs will have a minimal predatory impact on commercial R. philippinarum juvenile stocks.     

Ontogeny of Acoustic and Feeding Behaviour in the Grey Gurnard, Eutrigla gurnardus

Although sound production in teleost fish is often associated with territorial behaviour, little is known of fish acoustic behaviour in other agonistic contexts such as competitive feeding and how it changes during ontogeny. The grey gurnard, Eutrigla gurnardus, frequently emits knock and grunt sounds during competitive feeding and seems to adopt both contest and scramble tactics under defensible resource conditions. Here we examine, for the first time, the effect of fish size on sound production and agonistic behaviour during competitive feeding. We have made sound (alone) and video (synchronized image and sound) recordings of grey gurnards during competitive feeding interactions. Experimental fish ranged from small juveniles to large adults and were grouped in four size classes: 10–15, 15–20, 25–30 and 30–40 cm in total length. We show that, in this species, both sound production and feeding behaviour change with fish size. Sound production rate decreased in larger fish. Sound duration, pulse duration and the number of pulses increased whereas the peak frequency decreased with fish size, in both sound types (knocks and grunts). Interaction rate and the frequency of agonistic behaviour decreased with increasing fish size during competitive feeding sessions. The proportion of feeding interactions accompanied by sound production was similar in all size classes. However, the proportion of interactions accompanied by knocks (less aggressive sounds) and by grunts (more aggressive) increased and decreased with fish size, respectively. Taken together, these results suggest that smaller grey gurnards compete for food by contest tactics whereas larger specimens predominantly scramble for food, probably because body size gives an advantage in locating, capturing and handling prey. We further suggest that sounds emitted during feeding may potentially give information on the motivation and ability of the individual to compete for food resources.     

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.     

THE DIET, PREY SIZE AND CONSUMPTION OF PHILINE ORIENTALIS (OPISTHOBRANCHIA: PHILINIDAE) IN HONG KONG

Philine orientalis occurs on Hong Kong's beaches for {smalltilde}2 months each year, in May and June when it breeds, laysegg capsules and dies. It is believed to have a life span ofone year. Hoi Sing Wan, a sandy beach in the New Territoriesof Hong Kong is dominated at lower levels by the venerid bivalveTapes philippinarum, the population of which in May and June,following spring recruitment, mostly comprises juveniles. P.orientalis, occurring at a density of {small tilde} 1.m^–2,feeds voraciously upon these juveniles consuming, in laboratoryexperiments, a mean of 25.day^–1. At Hoi Sing Wan, P. orientalisappears to be highly prey specific. It is, however, known tofeed on other prey items and probably has a catholic diet, duringits annual migration from deeper to shallower sediments, accordingto the prevailing fauna. In terms of wet weight, P. orientalis consumes a mean of {smalltilde}4.4% (2.2–6.7%) of its body weight day^–1.In terms of dry weight, however, this figure increases to amean of 35.5% and one figure of 208% has been recorded. Althoughthis is because of the large body fluid content of this, essentiallyshell-less, gastropod, using a hydrostatic skeleton to achieveburrowing, such figures further attest to the voracity of thebreeding adults. This study joins others in questioning ourpresent understanding of the scale of energy budgets on suchshores, i.e., are our estimates of loss to predation grosslyunderestimated? ^*Present address: Department of Biology, Baptist College, WaterlooRoad, Kowloon, Hong Kong. (Received 20 May 1989; accepted 3 July 1989)     

The role of pelagic water mites in the control of cladoceran population in a temperate lake of the southern Andes

In a small temperate lake of the southern Andes, Bosmina longirostrisand Ceriodaphnia dubia coexist with the predaceous water miteLimnesia patagonica. Sampling of natural populations and laboratoryexperiments were carried out. The field population of Limnesiadid not show a numerical response to the density or biomassof its prey. Laboratory experiments showed that the water miterejected C.dubia adults and juveniles as prey, but consumedBosmina. The maximum predation rate was 40 prey predator^–1day^–1 and a linear relationship between predation rateand prey density was obtained (R² = 66%). The contribution ofmortality due to predation and the predation risk are too lowto provoke a prey suppression. By means of computer modelling,the densities of predator necessary to explain half of the totalprey mortality were calculated. These densities were one ortwo orders of magnitude higher than those in the field. It isconcluded that Limnesia could really be a suppressor, but thepotential depends greatly on its density.     

The effects of interspecific competition and prey odor on foraging behavior in the rock crab, Cancer irroratus (Say)

The effects of competitor pressure and prey odor on foraging behavior of the rock crab, Cancer irroratus (Say), were investigated. The Jonah crab, Cancer borealis (Stimpson), was chosen as the interspecific competitor because it shares resources with C. irroratus. Four treatments were tested for their effect on foraging: the presence or absence of a competitor and two types of prey odor; body odor (living mussel) and tissue extract (dead mussel tissue). The presence of Jonah crabs did not influence location time, search time, prey size selected, or handling time of the rock crabs. However, rock crabs responded differently to the presence of body odor and tissue extract cues. The presence of extract odor decreased the time to locate prey while increasing the number of prey manipulated and prey size selected. When prey body odor was present, rock crabs displayed less investigative behaviors than in the presence of extract odor, illustrated by reduced location time. Extract odor provided a stronger and more attractive cue than body odor, but increased prey manipulation and search time. Extract odor induced increases in manipulation and searching for prey but canceled out the benefits of decreased location time, resulting in crabs from both treatments displaying similar search times. These elevated behaviors may be associated with foraging for injured and cracked prey or may indicate an area of conspecific feeding.     

Interference competition,payoff asymmetries,and the social relationships of central place foragers

We present a central place foraging model that shows how payoff asymmetries originate in contests for access to resources. The essence of the model is that interference competition at resource points lowers the rate at which foragers can load prey, thereby depressing the rate of food delivery to the central place. We show that interference leads to asymmetric payoffs when contests involve foragers with (i) unequal travel distances between the central place and the contested resource points; (ii) inequalities in the rate of food delivery available from alternative foraging sites; (iii) differences in loading efficiency; or (iv) different abilities to interfere. We use the asymmetries to predict dominance rankings, and the patch exploitation tactics of individual foragers. We also consider the implications of the model for changes in the travel distance (= area) over which foragers can exclude competitors (= territoriality) as food density changes. Finally, incorporation of interference permits our model to predict the transition between scramble and contest competition.     

Balancing predation risk,social interference,and foraging opportunities in backswimmers,<Emphasis Type="Italic"> Notonecta maculata</Emphasis>

Loss of foraging opportunities and intraspecific competition for prey may be important costs of using refuges, because a hiding animal is unable to use or defend its foraging area from conspecific intrusions. Thus, animals should balance antipredator demands with other requirements in deciding when to come out from a refuge after a predators unsuccessful attack. Observations on foraging and social interactions of backswimmers Notonecta maculata suggest that foraging may be costly in terms of intraspecific agonistic interactions. When prey density is low, increasing the probability of finding a prey may require active exploration of a larger area, but this also increases the probability of encountering a competitor. After simulated exposure to predators, unfed bugs resumed feeding positions after a significantly shorter hiding period than recently fed bugs. We hypothesized that hiding time may also be reduced by recent interactions with conspecific competitors, due to an increased perceived need to defend feeding opportunities. Thus, when a predator attack occurred immediately after an agonistic conspecific interaction, backswimmers resumed feeding positions more quickly, and closer to the original position from which they were disturbed, suggesting short-term defense of particular positions. We conclude that when foraging, backswimmers balance the benefits of finding prey with the costs of predation risk and social interference in deciding their foraging strategy.Communicated by P.K. McGregor     

Kleptoparasitic prey competition in shoaling fish: effects of familiarity and prey distribution

Familiarity is thought to stabilize dominance hierarchies andreduce aggressive interactions within groups of socially livinganimals. Though familiarity has been widely studied in shoalingfish, few studies have investigated changes in prey competitionas a function of time spent together within groups of initiallyunfamiliar individuals. In this study, we created shoals ofthree-spined stickleback (Gasterosteus aculeatus) and monitoredchanges in foraging rates and related competitive behaviorswithin shoals over a 4-week period in experimental series whereprey was spatially and temporally concentrated or dispersed.Prey share was unequal under both prey distribution modes, anddisparity in prey share was not seen to change as trials progressed.Interestingly, the contest rate for prey items fell over timewhen individuals were competing for dispersed prey but not whenprey were concentrated. We found no evidence that fish showedassociation preferences for either group members that had consumeda greater or lesser proportion of prey during trials. Thoughthe intensity of competition may be reduced by increased groupstability in nature, this is likely to be strongly dependenton the way prey resources are distributed through space andtime.     

Cephalopods in the Diet of Albacore, Thunnus alalunga, from the Adriatic Sea

The stomachs of 56 albacore (Thunnus alalunga) caught in thesouthern Adriatic Sea contained 222 cephalopods, or 29% of prey.Most cephalopod remains were from small specimens. Juvenilesand adults of Heteroteuthis dispar (61.5% of cephalopods) were morefrequently preyed upon in the `inshore' fishing area (bottomdepth: 100-700 m); early juveniles of Todarodes sagittatus (21.6%)were more abundant ` offshore' (750- 1,150 m); early juvenilesof Histioteuthis bonnellii (16.4%) were exclusive of the latterarea. The estimated overall number of cephalopods dying of predationby albacore is 6.3 x 10⁶ specimens/fishing season (two months). (Received 6 May 1998; accepted 13 July 1998)     

Ontogenesis of agonistic vocalizations in the cichlid fish Metriaclima zebra

While acoustic communication has been described in adults of various fish species, our knowledge about the ontogeny of fish sound production is limited. In adults, sound signals are known to be involved during aggressive interactions. However, aggressive behaviour may appear early in the life of fishes due to the possible competition for food and space. If acoustic signals are used to send information to competitors, sounds are likely to play a role during interactions between juvenile fish as well. The apparition and evolution of sound production were monitored in a group of juveniles of the cichlid fish Metriaclima zebra from hatching to 4months of age. In addition, the link between vocalizations and agonistic behaviour was studied during dyadic interactions at three different ages. Sounds production appeared to be present early in the development of this fish and increased along with the number of aggressive behaviours. Recorded sounds consisted, in juveniles, in isolated pulses showing a decrease in frequency and duration as the fish grew. In adults, sounds became bursts of pulses but the transition from isolated to repetitive pulses was not observed. These results are compared to the existing literature on sound production ontogeny in fishes.     

Interspecific competition and coexistence between ants and land hermit crabs on small Bahamian islands

Numerous studies have demonstrated the existence of intra- and interspecific competition among ants, but few have investigated direct competitive interactions between ants and other taxa. In this paper, I present the first evidence of direct competitive interactions between ants and crabs. Evidence of competition for food between ants and the land hermit crab, Coenobita clypeatus (Herbst), was derived from observations and experiments in an archipelago of small islands in the central Exumas, Bahamas. Correlational evidence of competition for food based on occurrences at baits was found between ants and hermit crabs in multiple years. Observations at baits over time revealed species turnover occurred due to aggressive interactions. C. clypeatus discovered food items rapidly, but lost control of food over time, particularly to the ant Brachymyrmex obscurior Forel, which took longer to find food items but recruited large numbers of workers that drove off hermit crabs. A second ant species, Dorymyrmex pyramicus Roger, discovered baits quickly but did not recruit to baits in large numbers, and was not a superior competitor to either C. clypeatus or B. obscurior. Competition between ants and land hermit crabs was not intense enough to cause complementary distributions, and mechanisms of coexistence apparently include temporal variation in foraging activity and complementary foraging strategies when ants and crabs are active at the same time. Because of the widespread distributions and generalist scavenger diets of many ants and crabs, such competitive interactions are likely to be a common facet of many tropical and subtropical insular and coastal communities.     

Female and male Texas cichlids (Herichthys cyanoguttatum) do not fight by the same rules

Selection usually acts differently on males and females duringintrasexual competition for resources and/or mates. Nevertheless,agonistic behavior has been examined both theoretically andempirically mostly in males. Our research questions whethermales and females follow the same rules of engagement in intrasexualcontests as predicted by the sequential assessment model (SAM).The SAM predicts negative correlations between contest intensityand duration and the magnitude of asymmetry in resource holdingpower (RHP) between the contestants, such that the most escalatedcontests are those between similarly endowed individuals. Westaged male and female intrasexual contests with varying degreesof body size asymmetry under a round robin design using themonogamous Texas cichlid fish (Herichthys cyanoguttatum) asa study case. We used Mantel's matrix analysis to compare howthe behavioral content, duration, structure, and outcome ofmale and female contests were affected by the relative bodysize of the contestants. In the case of males, relative sizein each contest predicted outcome, duration, and frequency ofconventional and escalated behaviors according to prevailingtheory. Female contest structure and outcome, however, werenot predicted by the relative size of contestants. We discussour results in terms of other asymmetries that might be importantin structuring female contests, and we propose potential approachesto study female–female aggression.     

Harassment of adults by immatures in bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>): testing the Exploratory Aggression and Rank Improvement hypotheses

The immatures of many primate species frequently pester adult group members with aggressive behaviors referred to as a type of harassment. Although these behaviors are characteristic of immatures as they develop from infancy through adolescence, there have been few studies that specifically address the adaptive significance of harassment. Two functional hypotheses have been generated from observations of the behavior in chimpanzees. The Exploratory Aggression hypothesis describes harassment as a mechanism used by immatures to learn about the parameters of aggression and dominance behavior and to acquire information about novel, complex, or unpredictable relationships. The Rank Improvement hypothesis describes harassment as a mechanism of dominance acquisition used by immatures to outrank adults. This study investigated harassment of adults by immatures in a group of bonobos housed at the Columbus Zoo and compared the results to the predictions outlined by the Exploratory Aggression and Rank Improvement hypotheses. Although all immature bonobos in this group harassed adults, adolescents performed the behavior more frequently than did infants or juveniles and low-ranking adults were targeted more frequently than high-ranking. Targets responded more with agonistic behaviors than with neutral behaviors and the amount of harassment an individual received was significantly correlated with the amount of agonistic responses given. Furthermore, bouts of harassment were found to continue significantly more frequently when responses were agonistic than when they were neutral. Adolescents elicited mostly agonistic responses from targets whereas infants and juveniles received mostly neutral responses. These results support predictions from each hypothesis where harassment functions both as a mechanism of social exploration and as a tool to establish dominance rank.