红火蚁与热带火蚁间的干扰竞争

【目的】为了探讨入侵火蚁在我国成功定殖及其之间的竞争机制。【方法】运用行为学方法研究红火蚁Solenopsis invicta(Buren)和热带火蚁Solenopsis geminata(Fabricius)在个体水平和群体水平上的攻击性、攻击手段及合作能力。【结果】一对一攻击试验中,红火蚁和热带火蚁之间攻击级别多集中在3级,两种入侵蚂蚁间以相互威胁为主;红火蚁大型工蚁与热带火蚁兵、工蚁间最为好斗,其攻击级别达到4级的比例最高,分别为33.04%、37.92%。热带火蚁兵蚁与各型红火蚁间攻击强度差异不显著;热带火蚁工蚁与红火蚁小型工蚁之间的攻击性最强,其攻击性(3.49)显著高于热带火蚁工蚁与红火蚁大、中型工蚁的攻击性(3.32和2.97)。在攻击手段上,3级打斗时各型红火蚁更倾向以物理攻击主动威胁热带火蚁,而热带火蚁兵、工蚁会采取多种方式主动攻击红火蚁,双方皆以躲避应对为主;4级打斗时两种火蚁主要以混合攻击为主动或应对手段。群体攻击试验显示,红火蚁群体间攻击强度和合作性会随着群体数量的增加而显著增加,热带火蚁合作性较差,其群体对抗红火蚁的优势仅仅是由于个体数量的增加。【结论】红火蚁比热带火蚁具有更强的竞争优势。研究结果为入侵蚂蚁间不对称竞争机制和长期群落替代的内在原因提供理论基础。     

Selection on Arctic charr generated by competition from brown trout

We experimentally explored population‐ and individual‐level effects on Arctic charr (Salvelinus alpinus) resulting from resource competition with its common European competitor, the brown trout (Salmo trutta). At the population level, we compared performance of the two species in their natural sympatric state with that of Arctic charr in allopatry. At the individual level, we established selection gradients for morphological traits of Arctic charr in allopatric and in sympatric conditions. We found evidence for interspecific competition likely by interference at the population level when comparing differences in average performance between treatments. The growth and feeding rates did not differ significantly between allopatric and sympatric Arctic charr despite lower charr densities (substitutive design) in sympatric enclosures indicating that inter‐ and intraspecific competition are of similar strength. The two species showed distinct niche segregation in sympatry, and brown trout grew faster than Arctic charr. Arctic charr did not expand their niche in allopatry, indicating that the two species compete to a limited degree for the same resources and that interference may suppress the growth of charr in sympatric enclosures. At the individual level, however, we found directional selection in sympatric enclosures against individual Arctic charr with large head and long fins and against individuals feeding on zoobenthos rather than zooplankton indicating competition for common resources (possibly exploitative) between trout and these charr individuals. In allopatric enclosures these relations were not significant. Diets were correlated to the morphology supporting selection against the benthic‐feeding type, i.e. individuals with morphology and feeding behaviour most similar to their competitor, the benthic feeding brown trout. Thus, this study lends support to the hypothesis that Arctic charr have evolved in competition with brown trout, and through ecological character displacement adapted to their present niche.     

ALTERNATIVE PATHS TO SUCCESS IN A PARASITE COMMUNITY: WITHIN‐HOST COMPETITION CAN FAVOR HIGHER VIRULENCE OR DIRECT INTERFERENCE

Selection imposed by coinfection may vary with the mechanism of within‐host competition between parasites. Exploitative competition is predicted to favor more virulent parasites, whereas interference competition may result in lower virulence. Here, we examine whether exploitative or interference competition determines the outcome of competition between two nematode species (Steinernema spp.), which in combination with their bacterial symbionts (Xenorhabdus spp.), infect and kill insect hosts. Multiple isolates of each nematode species, carrying their naturally associated bacteria, were characterized by (1) the rate at which they killed insect hosts, and by (2) the ability of their bacteria to interfere with each other's growth via bacteriocidal toxins called “bacteriocins.” We found that both exploitative and interference abilities were important in predicting which species had a selective advantage in pairwise competition experiments. When nematodes carried bacteria that did not interact via bacteriocins, the faster killing isolate had a competitive advantage. Alternatively, nematodes could gain a competitive advantage when they carried bacteria able to inhibit the bacteria of their competitor. Thus, the combination of nematode/bacterial traits that led to competitive success depended on which isolates were paired, suggesting that variation in competitive interactions may be important for maintaining species diversity in this community.     

Foraging behavior as a determinant of asymmetric competitive interaction between two ant species in a tropical agroecosystem

This work is concerned with elucidating competitive interactions between two neotropical ants, Solenopsis geminata and Pheidole radoszkowskii, focusing on their foraging behavior. When released from competition from P. radoszkowskii, S. geminata increased its foraging activity. On the other hand, when released from competition from S. geminata, P. radoszkowskii did not respond, demonstrating asymmetric competition between the two species. Foraging experiments showed that P. radoszkowskii is more efficient at finding food resources, whereas S. geminata is better at defending the resources once they are encountered. These differences in foraging behavior appear to permit the coexistence of these two species. The practical implications of the results for the management of ant communities in tropical agroecosystems is discussed with respect to the potential use of ants as natural enemies.     

Mechanisms of interspecific competition among an invasive and two native fire ants

The mechanisms of interspecific competition among an invasive and two native Solenopsis fire ant forms were investigated in a series of laboratory experiments. In separate trials each with a different food resource, the native S. geminata × xyloni retrieved the greatest amount of a protein- and lipid-rich artificial food resource and a high protein natural food resource, and the native S. geminata retrieved the greatest amount of a high carbohydrate food resource. In trials investigating aspects of interference competition at the colony level, the invasive S. invicta proved to be initially more aggressive than S. geminata , but less aggressive than S. geminata × xyloni . Solenopsis invicta eventually controlled more of the foraging arenas against both native forms when colonies were equivalent by worker biomass, but not when colonies were equivalent by worker number. When paired with S. invicta , S. geminata suffered a significantly greater proportional reduction in both workers and entire colonies when colonies were initially standardized by worker biomass, but not when colonies were standardized by worker number. When paired with S. invicta , a significantly greater proportional reduction of workers occurred in S. geminata × xyloni , regardless of how colonies were standardized. In pairwise trials at the individual level, majors always exhibited significantly less mortality than minors, regardless of the Solenopsis form. The majors of both native forms suffered significantly less mortality than those of S. invicta . Superiority in colony-level interference ability appears to be an important mechanism allowing S. invicta to displace native Solenopsis forms. The ability of S. invicta to reach high population densities, because of intrinsic biological characteristics or an escape from natural enemies, plays an important contributory role. Similar mechanisms may underlie the success of other invasive ant species.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

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.     

Species richness,relative abundances and habitat use in local assemblages ofSorex shrews in Eurasian boreal forests

Using a large body of observational data on the occurrence ofSorex shrews in boreal forests, we test two models that predict the structure of small mammal communities along a gradient of increasing habitat productivity. Tilman’s (1982) model predicts a humped curve of species richness along productivity gradients. In contrast, we found a linear increase in species richness with increasing logarithm of the pooled density of shrews, which we use as a measure of habitat productivity for shrews. The model of Hanski and Kaikusalo (1989) assumes a trade-off between exploitative and interference competitive abilities, and it predicts that the size structure of small mammal communities should shift from the dominance of small species (superior in exploitative competition) in unproductive habitats to the dominance of large species (superior in interference competition) in productive habitats. Shrew assemblages show such a shift. Though it is not possible to draw definite conclusions about the role of interspecific competition from our observational data, the changing size structure of local shrew assemblages with increasing habitat productivity is a predictable feature of their community structure.     

Interference competition and species coexistence

Interference competition is ubiquitous in nature. Yet its effects on resource exploitation remain largely unexplored for species that compete for dynamic resources. Here, I present a model of exploitative and interference competition with explicit resource dynamics. The model incorporates both biotic and abiotic resources. It considers interference competition both in the classical sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, and interference on, the other species) and in the broad sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, but can experience an increase in growth rate via interference on, the other species). Coexistence cannot occur under classical interference competition even when the species inferior at resource exploitation is superior at interference. Such a trade-off can, however, change the mechanism of competitive exclusion from dominance by the superior resource exploiter to a priority effect. Now the inferior resource exploiter can exclude the superior resource exploiter provided it has a higher initial abundance. By contrast, when interference is beneficial to the interacting species, coexistence is possible via a trade-off between exploitation and interference. These results hold regardless of whether the resource is biotic or abiotic, indicating that the outcome of exploitative and interference competition does not depend on the exact nature of resource dynamics. The model makes two key predictions. First, species that engage in costly interference mechanisms (e.g. territoriality, overgrowth or undercutting, allelopathy and other forms of chemical competition) should not be able to coexist unless they also engage in beneficial interference mechanisms (e.g. predation or parasitism). Second, exotic invasive species that displace native biota should be superior resource exploiters that have strong interference effects on native species with little or negative cost. The first prediction provides a potential explanation for patterns observed in several natural systems, including plants, aquatic invertebrates and insects. The second prediction is supported by data on invasive plants and vertebrates.     

The Relative Role of Interference Competition in Regulation of a Rotifer Community during Spring Development in a Eutrophic Reservoir

The relative importance of interference and exploitative competition from Daphnia was tested in contrast to known role of Cyclops vicinus predation, influencing the rotifer community in Římov Reservoir during spring period. The abundances of five dominant rotifer species and their birth and death rates were analysed, together with changes in chl‐a concentration, HNF, bacteria biomass and the abundances of predator and competitor species, from high frequency sampling. Development of loricated species (Keratella cochlearis, K. quadrata and Kellicottia longispina) was driven predominantly by food availability. The dominant soft‐bodied species Polyarthra spp. and Synchaeta lakowitziana were affected predominantly through C. vicinus predation. The role of mechanical interference from D. galeata, although may be important in a limited periods was overclouded by predation and exploitative competition in general. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Costs of coexistence along a gradient of competitor densities: an experiment with arvicoline rodents

1. Costs of coexistence for species with indirect resource competition usually increase monotonically with competitor numbers. Very little is known though about the shape of the cost function for species with direct interference competition. 2. Here we report the results of an experiment with two vole species in artificial coexistence in large enclosures, where density of the dominant competitor species (Microtus agrestis) was manipulated. Experimental populations of the subordinate vole species (Clethrionomys glareolus) were composed of same aged individuals to study distribution of costs of coexistence with a dominant species within an age-cohort. 3. Survival and space use decreased gradually with increasing field vole numbers. Thus, responses to interference competition in our system appeared to be similar as expected from resource competition. The total number of breeders was stable. Reproductive characteristics such as the timing of breeding, and the litter size were not affected. In the single species enclosures a proportion of surviving individuals were not able to establish a breeding territory against stronger conspecifics. Under competition with heterospecifics such nonbreeders suffered high mortality, whereas the breeders survived. 4. Combined interference of dominant conspecifics and heterospecifics probably increased the frequency of aggressive interactions, social stress and mortality for the weaker individuals within a homogeneous age cohort of the subordinate competitor population. 5. Our results suggest, that in open systems where bank voles are outcompeted over the breeding season by faster reproducing field voles, animals able to establish a territory may be able to withstand competitor pressure, while nonbreeding bank vole individuals are forced to emigrate to suboptimal forest habitats.     

Alternatives to Lotka-Volterra competition: Models of intermediate complexity

A family of one-level differential-equation competition models in which two populations are limited by the energy flowing into the system generates the following results. For competitors on the same and only resource: 1) Purely exploitative competition leads to exclusion; which species wins depends on relative abilities to appropriate and extract energy from the resource, and the relative death and maintenance rates. 2) If conspecific interference (e.g., deaths or energy loss from fighting, cannibalism, or display) is sufficiently high relative to abilities to exploit the common resource, competition for the same resource can lead to coexistence. 3) If heterospecific interference is sufficiently high relative to abilities to exploit the common resource, competition for the same resource can lead to a priority effect, in which the outcome depends on initial population sizes. 4) Depending on whether situation (2) or (3) prevails, an increase in the amount of the common resource can convert an outcome in which one species always wins into one giving coexistence (2) or a priority effect (3). 5) If species are similar to one another in their abilities to appropriate and extract energy from the common resource and show reciprocity in intererence costs, competition can have multiple outcomes; either one species wins or the species coexist, depending on initial values.For competition on the same resource, but with each species monopolizing an exclusive resource as well: 1) Purely exploitative competition always leads to a unique point coexistence. 2) If interference is added to the system described in (1), two points of coexistence, separated by a saddle (an “unstable equilibrium”) are possible. This is favored by a) a small yield from the exclusive resources relative to the common one; and b) strong interspecific relative to intraspecific interference.     

Contrasting forms of competition set elevational range limits of species

How abiotic and biotic factors constrain distribution limits at the harsh and benign edges of species ranges is hotly debated, partly because macroecological experiments testing the proximate causes of distribution limits are scarce. It has long been recognized – at least since Darwin’s On the Origin of Species – that a harsh climate strengthens competition and thus sets species range limits. Using thorough field manipulations along a large elevation gradient, we show the mechanisms by which temperature determines competition type, resulting in a transition from interference to exploitative competition from the lower to the upper elevation limits in burying beetles (Nicrophorus nepalensis). This transition is an example of Darwin’s classic hypothesis that benign climates favor direct competition for highly accessible resources while harsh climates result in competition through resources of high rivalry. We propose that identifying the properties of these key resources will provide a more predictive framework to understand the interplay between biotic and abiotic factors in determining geographic range limits.     

Eavesdropping in solitary large carnivores: Black bears advance and vocalize toward cougar playbacks

Large carnivore behavioral responses to the cues of their competitors are rarely observed, but may mediate competition between these top predators. Playback experiments, currently limited to interactions involving group‐living large carnivores, demonstrate that attending to cues indicative of the immediate presence of heterospecific competitors plays a substantial role in influencing competition among these species. Group‐living species vocalize regularly to signal to one another, and competitors can readily “eavesdrop” on these acoustic cues. Solitary large carnivores also vocalize to conspecifics, but much less frequently, reducing the ease with which heterospecific competitors can eavesdrop. Eavesdropping could nonetheless play a substantive role in mediating competition among solitary large carnivores if the benefits of responding to the acoustic cues of heterospecific competitors (reducing risk or locating resources) are sufficiently large. Behavioral interactions between solitary large carnivore species are almost never observed, and there have been no experimental tests of their reactions to cues indicative of the immediate presence of other solitary large carnivores. We used an automated playback system to test the responses of a solitary large carnivore (black bear, Ursus americanus) to vocalizations of their similarly solitary competitor (cougar, Puma concolor), presenting both cougar and control vocalizations to free‐living bears foraging along shorelines in British Columbia, Canada. Both mothers with cubs and solitary bears were significantly more likely to advance and vocalize toward cougar than control playbacks, mothers producing one or both of two distinct vocalizations and solitary bears producing just one. Cougars could either represent a potential risk to bears (particularly cubs), or a source of resources, as bears are known to regularly scavenge cougar kills. Our results are consistent with bears eavesdropping on cougars for both these reasons. As with group‐living species, eavesdropping may be common among solitary large carnivores, and may be an important driver of competition between these species.     

Phorid fly parasitoids of invasive fire ants indirectly improve the competitive ability of a native ant

Abstract.  1. The red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), is an invasive species of south-eastern U.S.A. Since its introduction from South America approximately 70 years ago, this pest has devastated natural biodiversity.
2. Due to such ecological costs, Pseudacteon phorid fly parasitoids (Diptera: Phoridae) from South America are being introduced into the U.S.A. as a potential biological control agent. Here, the indirect effects of these specialised parasitoids on an interspecific native ant competitor, Forelius mccooki (Hymenoptera: Formicidae), are evaluated.
3. Over the course of a 50-day laboratory experiment, the results show that the native ant improved aspects of exploitative, but not interference, competition when S. invicta -attacking flies were present compared with when they were absent.
4.  Forelius mccooki colonies from the phorid treatment had approximately twice as many foragers at food baits relative to controls; however, there was no significant difference in interference aspects of competition or native ant colony growth between the two treatments.
5. These results suggest that the S. invicta -specialised parasitoids help shift the competitive balance more in favour of F. mccooki than if these flies were not present; however, this competitive advantage does not translate into increased colony growth after 50 days. These laboratory findings are interpreted with regard to the more complex interactions in the field.     

Contest and scramble competitions in two bruchid species,Callosobruchus analis andC. phaseoli (Coleoptera: Bruchidae)

Larval competition between contest and scramble strategists was investigated using the two bruchid species, C. analis (contest species) and C. phaseoli (scramble species) with two different sized mung beans (large and small beans). In both sized beans, the adult emergences of each species dependen on total density of the initial larval densities of the two species and the ratio of the two densities. The emergence of one species was suppressed by the existence of the other species when the initial larval density per bean of the former species was less than that of the latter one. There were many cases in which both C. analis and C. phaseoli emerged from one bean in large beans, but such cases were quite rare in small beans. C. analis performed interference behavior only at late larval stages, whereas C. phaseoli was superior in exploitative competition all through their larval stages. These, combined with the niche segregation inside a bean, are throught to be the major factors of observed density- and frequency-dependent competition results. Based on the above experimental results, long-term competition results between the contest and scramble species were predicted.     

Effect of scavenging on predation in a food web

Scavenging can have important consequences for food web dynamics, for example, it may support additional consumer species and affect predation on live prey. Still, few food web models include scavenging. We develop a dynamic model that includes two facultative scavenger species, which we refer to as the predator or scavenger species according to their natural scavenging propensity, as well as live prey, and a carrion pool to show ramifications of scavenging for predation in simple food webs. Our modeling suggests that the presence of scavengers can both increase and decrease predator kill rates and overall predation in model food webs and the impact varies (in magnitude and direction) with context. In particular, we explore the impact of the amount of dynamics (exploitative competition) allowed in the predator, scavenger, and prey populations as well as the direction and magnitude of interference competition between predators and scavengers. One fundamental prediction is that scavengers most likely increase predator kill rates, especially if there are exploitative feedback effects on the prey or carrion resources like is normally observed in natural systems. Scavengers only have minimal effects on predator kill rate when predator, scavenger, and prey abundances are kept constant by management. In such controlled systems, interference competition can greatly affect the interactions in contrast to more natural systems, with an increase in interference competition leading to a decrease in predator kill rate. Our study adds to studies that show that the presence of predators affects scavenger behavior, vital rates, and food web structure, by showing that scavengers impact predator kill rates through multiple mechanisms, and therefore indicating that scavenging and predation patterns are tightly intertwined. We provide a road map to the different theoretical outcomes and their support from different empirical studies on vertebrate guilds to provide guidance in wildlife management.     

Dung beetle assemblages in forests and pastures of El Salvador: a functional comparison

This study examines the functional attributes of Neotropical dung beetles, and, based on the analysis, evaluates the role of dung beetles in pastures and tropical dry forest in El Salvador. Dung beetle diversity was lower in pastures than in forests. However, the total biomass of beetles at similarly-sized dung baits was frequently higher in pastures. Diversity loss followed structured patterns: (1) carrion and fruit feeding beetles were well represented (species number) in the forests but were largely absent from pastures; (2) large ball rollers and small fast tunnellers were present in forests but were absent from pastures; (3) large fast tunnellers and small slow tunnellers were poorly represented in pastures compared to forests but the few species that survived in pastures attained extremely high populations; and (4) each functional group in the species-poor pasture assemblages was divided almost equally into day and night active species. Substrate and habitat generalists that were present in both the forests and pastures were attracted to fruit and carrion in forested habitat but not in open pastures. In open habitats, flies and other insects may be the principal decomposers of decaying fruit and carrion.     

On a resource based ecological competition model with interference

A resource based ecological competition model with interference is proposed. The model is based on Lotka-Volterra dynamics with two predators competing for a single, limited prey. Interference effects are considered in this article. When the interference coefficient, expressing the damage effect from its rival, is small, the mathematical analysis shows that the winner in purely exploitative competition still outcompetes its rival. However, if the interference coefficient is large enough then the competition outcome will depend on initial population of predator species.     

Interference from a game theoretical perspective: shore crabs suffer most from equal competitors

In virtually all natural systems, interference competition amongindividuals is asymmetric. Here, we used game theoretical modelson contest behavior to predict how time lost in agonistic interactionscould affect strength of interference under asymmetric competition.We hypothesized that interference through time lost in agonisticinteractions would result in a greater reduction in availableforaging time and overall feeding rate under symmetric competitionthan under asymmetric competition. We tested this hypothesisfor male shore crabs (Carcinus maenas) that foraged on mussels(Mytilus edulis) in an experiment where prey levels were keptconstant. We varied absolute size of crabs (juveniles, smalladults, large adults), mussel density (4, 16, 32 per 0.25 m²),and competitor size (smaller, equal, larger). Large adults spentmore time in aggressive behaviors than juveniles or small adults,possibly because large adults were more persistent in interferingor because large adults were intrinsically more aggressive,as the experiment was conducted in the mating season. When handlingprey, crabs mostly avoided competitors, but juveniles and smalladults did so more than large adults. When searching for prey,crabs mostly displaced smaller competitors but threatened oravoided size-matched or larger competitors. By avoiding a competitor,the focal crab lost time but the competitor often did not, andthis asymmetry in agonistic behavior is not yet incorporatedin models on contest behavior. However, overall, negative effectsof others were strongest with size-matched competitors, in linewith our hypothesis.