Exploitative Competition Research Articles

Examining spatial patterns of selection and use for an altered predator guild.

Anthropogenic disturbances have altered species' distributions potentially impacting interspecific interactions. Interference competition is when one species denies a competing species access to a resource. One mechanism of interference competition is aggression, which can result in altered space-use of a subordinate species due to the threat of harm, otherwise known as a 'landscape of fear'. Alternatively, subordinates might outcompete dominant species in resource-poor environments via a superior ability to extract resources. Our goal was to evaluate spatial predictions of the 'landscape of fear' hypothesis for a carnivore guild in Newfoundland, Canada, where coyotes recently immigrated. Native Newfoundland carnivores include red foxes, Canada lynx, and black bears. We predicted foxes and lynx would avoid coyotes because of their larger size and similar dietary niches. We used scat-detecting dogs and genetic techniques to locate and identify predator scats. We then built resource selection functions and tested for avoidance by incorporating predicted values of selection for the alternative species into the best supported models of each species. We found multiple negative relationships, but notably did not find avoidance by foxes of areas selected by coyotes. While we did find that lynx avoided coyotes, we also found a reciprocal relationship. The observed patterns suggest spatial partitioning and not coyote avoidance, although avoidance could still be occurring at different spatial or temporal scales. Furthermore, Newfoundland's harsh climate and poor soils may swing the pendulum of interspecific interactions from interference competition to exploitative competition, where subordinates outcompete dominant competitors through a superior ability to extract resources.

Larval interference competition between the native Neotropical mosquito Limatus durhamii and the invasive Aedes aegypti improves the fitness of both species.

Interspecific competition with native species during biological invasions can sometimes limit alien expansion. We aimed to determine the potential ecological effects of Limatus durhamii Theobald 1901, a native Neotropical mosquito (Diptera: Culicidae) species, on the invasive species Aedes (Stegomyia) aegypti (Linnaeus 1762) that breeds in the same artificial water containers. Development time and adult dry mass were measured in 3 rearing conditions: control (a single larva), intraspecific competition (2 conspecific larvae), and interspecific competition (2 heterospecific larvae). Food was provided ad libitum to eliminate exploitative competition. For Ae. aegypti, development time was not affected by interspecific interference competition (nonsignificant differences with the control) and the adult dry mass was significantly higher, meaning that individual fitness likely increased. Yet, because previous studies showed longer development time and lighter adults during competition with other invasive mosquitoes, it is likely that Ae. aegypti can express a different phenotype depending on the competing species. The similar pattern found for Li. durhamii females and the nonsignificant difference with the control for males explain in part why this species can compete with Ae. aegypti.

Size and variation in individual growth rates among food web modules

AbstractThe complexity of food webs can be reduced to fundamental modules of trophic interactions that repeat to form reticulate webs. Omnivory, defined as feeding on more than one trophic level, is a module that is found in food webs more often than predicted by chance, likely because it confers stability. Yet, little is known about how omnivore and other food web modules affect individuals in the food web. Here, we constructed four different experimental food web modules using a blue mussel predator, zooplankton consumer, and phytoplankton resource. By manipulating the predator's access to the consumer and providing resource subsidies, we produced exploitative competition, food chain and omnivory food web modules, and a consumer‐resource interaction. We used RNA:DNA ratios to measure the growth rate of the consumer in each food web. In 24‐h experiments, growth rates of the consumer in the omnivory food web were significantly higher and more variable than in the other modules. Our results suggest that higher growth rates and variation at the individual scale may weaken the strength of predator–consumer interactions and help explain the ubiquity of omnivory in real food webs.

Effects of competition and climate on a crop pollinator community

Abstract Plant-pollinator interactions are ubiquitous in nature where both wild and domesticated pollinators interact with wild plant communities and entomophilous crops. Honeybees are important pollinators in many crop systems, but recent declines in honeybee stocks in Europe and the US have caused concern about the sustainability of crop systems solely depending on honeybees. In addition, several studies have shown that honeybees might negatively affect native pollinator populations, bumblebees in particular. Here we have studied flower visitation to two raspberry farms and surrounding wildflower communities in SE Norway. Bumblebees were excluded from the raspberry field by means of exploitative competition from honeybees ( > 97% of flower visits in the raspberry fields were conducted by honeybees). More than 55% of the visits recorded in wild plant communities surrounding the farms were conducted by bumblebees, showing that bumblebees were present in the system. Pollinator taxa were affected differently by temperature; honeybee visits showed a unimodal relationship with maximum flower visitation activity at a temperature of 24.1 °C, while flower visits by bumblebees showed a positive, linear relationship with temperature. The effect of temperature was much weaker for bumblebees than for honeybees (∼2.2% of the variation was explained by temperature, compared to ∼46% for honeybees). Farming practice affected flower visitation, as flowers within growing tunnels received fewer visits. However, the number of flower visits, also within the growing tunnels, was far above what other studies have shown to be sufficient for optimal pollination in raspberry. We conclude that the raspberry fields were sufficiently pollinated by honeybees but that the system should be considered vulnerable as it is solely dependent on this particular pollinator species. The honeybees were sensitive to ambient temperature suggesting that they might suffer more from future climate change than bumblebees.

Invasive ant (Anoplolepis gracilipes) disrupts pollination in pumpkin

Yellow crazy ant (Anoplolepis gracilipes (F. Smith); “YCA”) is known for its aggressive predatory ability and ability to exert exploitation competition on both native and other invasive ants via floral nectar. We argue that YCA invasion can exert both interference and exploitation competition on legitimate pollinators. In pumpkin fields (Cucurbita maxima L.) of south India, YCA infested the flowers, particularly the pistillate flowers, for nectar foraging. Pumpkin is a honey bee-mediated cross-pollinated monoecious plant that produces disproportionately very few pistillate flowers. We hypothesize that YCA presence in the flowers can affect the visitation rate and foraging time of honey bees in the flowers, the fruit set in pumpkins, and can exert predatory pressure on the honey bees if the bees linger in ant-colonized flowers. Both YCA and honey bees preferred to forage on the limited pistillate flowers in the plants. After colonizing the flowers, YCA did not retreat for hours, even upon disturbance by competitors, such as honey bees. Both the visitation frequency and the foraging time of honey bees were drastically reduced in ant-colonized flowers, and none of the ant-colonized flowers developed into fruits, suggesting that the YCA exert both an ecological and evolutionary pressure on pumpkin. The ants preyed upon about 17% of the honey bees that lingered in ant-colonized flowers, and the time the bees spent foraging predicted the fate of the bees. Exploitation competition exerted by the YCA on pumpkin may have far-reaching consequences for the pollination and productivity of this cash crop.

A window of opportunity: Subdominant predators can use suboptimal prey.

Introduced species have been linked to declines of native species through mechanisms including intraguild predation and exploitative competition. However, coexistence among species may be promoted by niche partitioning if native species can use resources that the invasive species cannot. Previous research has shown that some strains of the aphid Aphis craccivora are toxic to a competitively dominant invasive lady beetle, Harmonia axyridis. Our objective was to investigate whether these aphids might be an exploitable resource for other, subdominant, lady beetle species. We compared larval development rate, survival, and adult weight of five lady beetle species in no‐choice experiments with two different strains of A. craccivora, one of which is toxic to H. axyridis and one that is nontoxic. Two lady beetle species, Cycloneda munda and Coleomegilla maculata, were able to complete larval development when feeding on the aphid strain that is toxic to H. axyridis, experiencing only slight developmental delays relative to beetles feeding on the other aphid strain. One species, Coccinella septempunctata, also was able to complete larval development, but experienced a slight reduction in adult weight. The other two lady beetle species, Hippodamia convergens and Anatis labiculata, demonstrated generally low survivorship when consuming A. craccivora, regardless of aphid strain. All five species showed increased survival and/or development relative to H. axyridis on the “toxic” aphid strain. Our results suggest that this toxic trait may act as a narrow‐spectrum defense for the aphids, providing protection against only some lady beetle enemies. For other less‐susceptible lady beetles, these aphids have the potential to provide competitive release from the otherwise dominant H. axyridis.

How to evaluate the potential occurrence of intraguild predation

Intraguild predation is the combination of exploitative competition and predation among potential competitors that use similar resources. It has the potential to shape population dynamics and community structure. Although there is much empirical evidence for the occurrence of intraguild predation in natural ecosystems, the study of its effects is mainly limited to short-term microcosm experiments. There is, therefore, certain skepticism about its actual significance in nature. A relevant concern is that there is no consensus regarding criteria to evaluate the possible occurrence of intraguild predation in short-term experiments, and methodological differences may therefore underlie apparent inconsistencies among studies. Our purpose here was to evaluate existing criteria to offer guidance for the design of experiments to determine whether two species may potentially engage in intraguild predation. The criteria are based on the condition that intraguild predators need to experience immediate energetic gains when feeding on the intraguild prey. Thus, a relevant experimental design must quantify predation but also fitness benefits of feeding on the other species, i.e. increases in reproduction, somatic growth, or survival.

Is bigger really better? Relative and absolute body size influence individual growth rate under competition.

Models suggest that the mechanism of competition can influence the growth advantage associated with being large (in absolute body size or relative to other individuals in the population). Large size is advantageous under interference, but disadvantageous under exploitative competition. We addressed this prediction in a laboratory experiment on Rana temporaria tadpoles competing for limited food. There were 166 target individuals spanning a 10‐fold range in body mass reared for 3 days with three other individuals that were either the same size, half as large, or twice as large as the target. Relative growth rate (proportion per day) declined with size, and absolute growth rate (mass per day) reached a peak at intermediate size and declined thereafter. Tadpoles grew slowly if they were large relative to their competitors, although relative body size was less important than absolute size. As a result, size variation declined in groups that were initially composed of individuals of variable size. Thus, bigger was not better under exploitative competition. Our results help connect individual‐level behavior with individual growth and the size distribution of the population.

A temporal race-for-fish: The interplay between local hotspots of flatfish and exploitation competition between beam trawlers after a seasonal spawning closure

In this paper we examine the response of the Belgian beam trawl fishery after the implementation of a spawning closure in the Celtic Sea. It was observed that fishing effort was mainly reallocated in time, resulting in a short-term “race for fish” immediately after re-opening of the fishery. The rationale of this behavior was examined by analyzing the landing rate of the target species (sole) in a generalized additive mixed model. Results showed that daily sole landings were up to twice as high just after re-opening of the fishery and dropped within 3 weeks to a reference level. This pattern of landing rate per unit (lpue) of sole was strongly related to the spatial distribution of fishing effort in the closed area during the month of re-opening. During the first week, fishing effort was very patchy, with increasing dispersal toward the end of the month. These results give indirect evidence for the occurrence of high local concentration of sole as a result of the closure, and the occurrence of exploitation competition when the fishery restarted.

Predation risk influences feeding rates but competition structures space use for a common Pacific parrotfish.

In terrestrial systems it is well known that the spatial patterns of grazing by herbivores can influence the structure of primary producer communities. On coral reefs, the consequences of varied space use by herbivores on benthic community structure are not well understood, nor are the relative influences of bottom-up (resource abundance and quality), horizontal (competition), and top-down (predation risk) factors in affecting spatial foraging behaviors of mobile herbivorous fishes. In the current study we quantified space use and feeding rates of the parrotfish, Chlorurus spilurus, across a strong gradient of food resources and predator and competitor abundance across two islands with drastically different fisheries management schemes. We found evidence that while feeding rates of this species are affected by direct interference competition and chronic predation risk, space use appears to be primarily related to exploitative competition with the surrounding herbivore community. We found no evidence that predation risk influences diurnal foraging space use in this small bodied parrotfish species. Additionally, we found the influence of chronic predation risk on feeding rates of this species to be less dramatic than the results of recent studies that used model predators to measure acute behavioral responses of other species of herbivorous fishes. Our results indicate that the non-consumptive effects of predators on the foraging behaviors of coral reef herbivores may be less dramatic than previously thought.

Consumptive and nonconsumptive effect ratios depend on interaction between plant quality and hunting behavior of omnivorous predators

SummaryPredators not only consume prey but exert nonconsumptive effects in form of scaring, consequently disturbing feeding or reproduction. However, how alternative food sources and hunting mode interactively affect consumptive and nonconsumptive effects with implications for prey fitness have not been addressed, impending functional understanding of such tritrophic interactions. With a herbivorous beetle, two omnivorous predatory bugs (plant sap as alternative food, contrasting hunting modes), and four willow genotypes (contrasting suitability for beetle/omnivore), we investigated direct and indirect effects of plant quality on the beetles key reproductive traits (oviposition rate, clutch size). Using combinations of either or both omnivores on different plant genotypes, we calculated the contribution of consumptive (eggs predated) and nonconsumptive (fewer eggs laid) effect on beetle fitness, including a prey density‐independent measure (c:nc ratio). We found that larger clutches increase egg survival in presence of the omnivore not immediately consuming all eggs. However, rather than lowering mean, the beetles generally responded with a frequency shift toward smaller clutches. However, female beetles decreased mean and changed clutch size frequency with decreasing plant quality, therefore reducing intraspecific exploitative competition among larvae. More importantly, variation in host plant quality (to omnivore) led to nonconsumptive effects between one‐third and twice as strong as the consumptive effects. Increased egg consumption on plants less suitable to the omnivore may therefore be accompanied by less searching and disturbing the beetle, representing a “cost” to the indirect plant defense in the form of a lower nonconsumptive effect. Many predators are omnivores and altering c:nc ratios (with egg retention as the most direct link to prey fitness) via plant quality and hunting behavior should be fundamental to advance ecological theory and applications. Furthermore, exploring modulation of fitness traits by bottom‐up and top‐down effects will help to explain how and why species aggregate.

When pork is not on the menu: Assessing trophic competition between large carnivores and poachers

Overexploitation of wildlife for meat is a widespread phenomenon, which drives populations of many species toward extinction and may in turn affect large carnivores. Therefore, human hunters may compete with large carnivores over food resources and threaten their survival. In this study, we assessed the trophic competition of endangered Persian leopard with local poachers in Golestan National Park, Iran, where poaching has depleted populations of three ungulate species by 66–89% in the past decades. We compared leopard diet (77 scats) with prey offtake by poachers (75 poacher seizure records). In addition, we estimated prey abundance by line transect sampling (186km), camera trapping (2777 camera days), double-observer point-counts (64 scans) and dung counts (38km). Using interview surveys with local poachers, we also quantified their stated hunting preference. We documented a narrow hunting specialization of leopard (niche breadth 0.24) and poachers (niche breadth 0.19), and exclusivity (niche overlap 0.31) of their dietary/hunting niches, which suggest no exploitative competition between these two apex predators. This pattern likely results from the major role of wild boar in leopard diet. Due to religious beliefs, poachers avoid hunting this species and its population has increased in contrast to other ungulates. Considering the general avoidance of Suidae species across leopard range, depletion of alternative prey species may have resulted in a prey-switching strategy by leopard. The influence of religious beliefs and taboos on hunting preference and, consequently, on prey populations and predators' trophic niches shows the importance of incorporation of cultural beliefs in conservation practices.

Asymmetric competition for nectar between a large nectar thief and a small pollinator: an energetic point of view.

There are two alternative hypotheses related to body size and competition for restricted food sources. The first one supposes that larger animals are superior competitors because of their increased feeding abilities, whereas the second one assumes superiority of smaller animals because of their lower food requirements. We examined the relationship between two unrelated species of different size, drinking technique, energy requirements and roles in plant pollination system, to reveal the features of their competitive interaction and mechanisms enabling their co-existence while utilising the same nectar source. We observed diurnal feeding behaviour of the main pollinator, the carpenter bee Xylocopa caffra and a nectar thief, the northern double-collared sunbird Cinnyris reichenowi on 19 clumps of Hypoestes aristata (Acanthaceae) in Bamenda Highlands, Cameroon. For comparative purpose, we established a simplistic model of daily energy expenditure and daily energy intake by both visitor species assuming that they spend all available daytime feeding on H. aristata. We revealed the energetic gain-expenditure balance of the studied visitor species in relation to diurnal changes in nectar quality and quantity. In general, smaller energy requirements and related ability to utilise smaller resources made the main pollinator X. caffra competitively superior to the larger nectar thief C. reichenowi. Nevertheless, sunbirds are endowed with several mechanisms to reduce asymmetry in exploitative competition, such as the use of nectar resources in times of the day when rivals are inactive, aggressive attacks on carpenter bees while defending the nectar plants, and higher speed of nectar consumption.

Foreign Higher Education in India: The Latest Developments

India has long had mixed emotions about letting foreign higher education institutions operate in the country. Advocates champion greater student choice and high quality programming while opponents worry about exploitation and unfair competition. In practice, many foreign institutions do operate but the rules are complex and unevenly enforced. The latest government measures are more open and clear in some respects but otherwise perpetuate restrictions and ambiguity.

Seasonal niche segregation of two sympatric xenocyprinid fishes

Xenocypris microlepis (Bleeker, 1871) and Xenocypris davidi (Bleeker, 1871) are two closely related xenocyprinid species potentially subject to exploitative competition. They often coexist in many freshwater ecosystems, which offers an opportunity to analyze their relationships. The present study describes the seasonal feeding habits of age‐3 X. microlepis and X. davidi in Xin'anjiang Reservoir, located in Zhejiang, China, as determined by stable isotope analysis. Particulate organic matter (POM; 50%) and periphyton (50%) were the major items observed in the X. microlepis diet, whereas periphyton (>60%) was prevalent in the X. davidi diet. Seasonal variations were observed in the diet compositions of both xenocyprinid species. Whereas X. microlepis tended to ingest more sediment organic matter (SOM) in autumn than in spring and summer, SOM was the secondary food item for X. davidi over the three seasons. The seasonal diet shift observed in this work indicates niche segregation of the sympatric pair. The results of this study contribute to the current knowledge of seasonal trophic interactions in predator–prey systems where sympatric xenocyprinids are present.

Competitors and predators alter settlement patterns and reproductive success of an intraguild prey

AbstractThe spatial distribution of predators is affected by intra‐ and interspecific interactions within the predator guild. Studying these interactions under fluctuating food availability, while taking habitat characteristics into account, offers a quasi‐experimental setup to determine the relative impact of con‐ and heterospecifics on reproductive success of predators. We analyzed the settlement decisions and reproductive success of Eurasian Pygmy Owls (Glaucidium passerinum) in the presence of both breeding conspecifics and their competitor and intraguild predator, Tengmalm's Owls (Aegolius funereus), under fluctuating abundance of their primary prey (voles). We used detailed data, collected across 11 years in a large study area (1300 km²), on the spatial and temporal variation of habitat characteristics in all available nesting sites, local densities of both species, and vole abundance. We found that Pygmy Owls strongly avoided breeding close to conspecifics but did not avoid Tengmalm's Owl nests. Nest box occupation of Pygmy Owls was positively correlated to the proportion of old spruce, mature and old pine forests, and farmlands, and occupation probability was higher at high vole abundance and in areas occupied in previous years. Pygmy Owl avoidance of conspecifics decreased when voles were abundant. At high conspecific density, Pygmy Owls showed lower hatching success and delayed hatching date, in agreement with the observed spatial avoidance. Finally, even though breeding Pygmy Owls did not spatially avoid Tengmalm's Owls, the density of heterospecifics correlated with low hatching and fledging success in Pygmy Owls. This suggests short term fitness costs when living close to competitors, even when lethal effects of intraguild interactions are subtle. Our results suggest that joint costs of exploitative and interference competition by Tengmalm's Owls, as well as intraguild predation, were lower than those induced by intraspecific competition only. This result might be due to the similar body size between the species considered, Tengmalm's Owls being only twice as large as Pygmy Owls. Interactions among con‐ and heterospecifics can therefore modify the spatial settlement and reproductive success of individuals on a landscape scale, also within the predator guild.

The Impacts of Crustacean Zooplankton on a Natural Ciliate Community: a Short-term Incubation Experiment

Direct and indirect effects of crustacean zooplankton (cladocerans and copepods) are important regulators of ciliate communities, especially in eutrophic systems. However, it is not clear whether pseudodiaptomids (e.g., Schmackeria), one of the dominant calanoid copepods in Chinese lakes, effectively impacts natural ciliate communities. The impacts of small-bodied cladocerans (e.g., Bosmina) on ciliates are also controversial. We performed an incubation experiment using winter lake water from Lake Chaohu to assess the structuring effects that crustacean zooplankton have on natural ciliate populations. The presence and absence of cladocerans (Bosmina sp.) and copepods (Schmackeria inopinus) were alternated in four treatments. Both Bosmina sp. and Schmackeria inopinus had substantial impacts on ciliate abundance, biomass, and community structure. The response of ciliates was different in the presence of Bosmina sp. compared with Schmackeria inopinus and varied among categories such as the ciliate population, relative body size and functional feeding group. Our results also highlight the importance of interference and exploitative competition among metazooplankton groups.

Diet patterns of island foxes on San Nicolas Island relative to feral cat removal

Island foxes (Urocyon littoralis) are a species of conservation concern that occur on six of the Channel Islands off the coast of southern California. We analysed island fox diet on San Nicolas Island during 2006–12 to assess the influence of the removal of feral cats (Felis catus) on the food use by foxes. Our objective was to determine whether fox diet patterns shifted in response to the cat removal conducted during 2009–10, thus indicating that cats were competing with foxes for food items. We also examined the influence of annual precipitation patterns and fox abundance on fox diet. On the basis of an analysis of 1975 fox scats, use of vertebrate prey – deer mice (Peromyscus maniculatus), birds, and lizards – increased significantly during and after the complete removal of cats (n = 66) from the island. Deer mouse abundance increased markedly during and after cat removal and use of mice by foxes was significantly related to mouse abundance. The increase in mice and shift in item use by the foxes was consistent with a reduction in exploitative competition associated with the cat removal. However, fox abundance declined markedly coincident with the removal of cats and deer mouse abundance was negatively related to fox numbers. Also, annual precipitation increased markedly during and after cat removal and deer mouse abundance closely tracked precipitation. Thus, our results indicate that other confounding factors, particularly precipitation, may have had a greater influence on fox diet patterns.

Survival and Predation of Weasels (Mustela erminea, Mustela frenata) in North America

Predators may limit populations of small carnivores indirectly through exploitative competition for resources, and directly through interference competition, including predation. We evaluated direct effects of predation by large predators on small, specialist carnivores by estimating annual survival and cause-specific mortality. We used known fate models to estimate annual survival of 23 radio-collared short-tailed weasels (Mustela erminea) in western Oregon in 2012–2013. Further, we reviewed published and unpublished studies of radio-tracked weasels (Mustela spp.) in North America, to identify main predators of weasels and survival. If the study did not report survival but did report number of days tracked and causespecific mortalities, we extrapolated an annual survival index, based on daily survival rates (Mayfield index), to facilitate comparisons between studies. Estimated mean ± SD annual survival of short-tailed weasels in our study was 0.26 ± 0.08, and did not vary by season or sex of the weasel. Most (80%) predation mortalities, however, occurred in winter, and were attributed to predatory birds, suspected to be barred owls (Strix varia). For seven North American studies, including ours, we estimated the average annual survival index for M. erminea and Mustela frenata to be 0.37 ± 0.28. Across studies, most mortality was attributed to predation. Predators killed weasels > 2x more frequently in winter when most mortality (86%) was attributed to predatory birds. Mammalian predators, in contrast, killed weasels most frequently in summer (75%). Our results support the hypothesis that predation may play an important role limiting weasel populations in North America.

Modulation of the gut microbiota by prebiotic fibres and bacteriocins.

ABSTRACTThe gut microbiota is considered an organ that co-develops with the host throughout its life. The composition and metabolic activities of the gut microbiota are subject to a complex interplay between the host genetics and environmental factors, such as lifestyle, diet, stress and antimicrobials. It is evident that certain prebiotics, and antimicrobials produced by lactic acid bacteria (LAB), can shape the composition of the gut microbiota and its metabolic activities to promote host health and/or prevent diseases. In this review, we aim to give an overview of the impact of prebiotic fibres, and bacteriocins from LAB, on the gut microbiota and its activities, which affect the physiology and health of the host. These represent two different mechanisms in modulating the gut microbiota, the first involving exploitative competition by which the growth of beneficial bacteria is promoted and the latter involving interference competition by which the growth of pathogens and other unwanted bacteria is prevented. For interference competition in the gut, bacteriocins offer special advantages over traditional antibiotics, in that they can be designed to act towards specific unwanted bacteria and other pathogens, without any remarkable collateral effects on beneficial microbes sharing the same niche.