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Competition between tadpoles and mosquito larvae.

Tadpoles and mosquito larvae often co-occur, and may compete for scarce resources. However, competition between such distantly related organisms has attracted less scientific attention than have interactions among closely related taxa. We examined ecological interactions in two tadpole-mosquito systems in southeastern Australia, one from freshwater ponds (Limnodynastes peronii and Culex quinquefasciatus) and one from brackish-water habitats (Crinia signifera and Ochlerotatus australis). Diets of these tadpoles and mosquito larvae overlap considerably, potentially leading to competition for food. Laboratory experiments show that, in both study systems, mosquitoes reduced the growth rates of tadpoles, and tadpoles reduced the growth rates and survival of mosquito larvae. These negative effects were seen even at high food levels. Thus, our study suggests that tadpoles and mosquito larvae affect each other strongly, and do so via pathways other than simple consumptive competition. Because mosquitoes are important vectors for human diseases, the global decline in amphibian populations may have more impact on human health than has generally been anticipated.

Analysis of Variance↗

Sympatry and allopatry in two desert ant sister species: how do Cataglyphis bicolor and C. savignyi coexist?

Two extremely morphologically similar sister species of desert ants, Cataglyphis bicolor and C. savignyi, exhibit broadly overlapping distributional ranges within Tunisia. In order to analyse the microhabitats of C. bicolor and C. savignyi within the sympatric and allopatric areas of both ant species, the plant species located at 113 different nest sites of the two ant species were determined. In the sympatric area, the two species exhibit a clear-cut nest site segregation. This is not the case in the allopatric areas. Hence the two species differentiate their microhabitat only when they are sympatric. The plant species associated mainly with the nest sites of C. bicolor indicate that this species prefers a type of vegetation that needs irrigation. This is in contrast to the nest sites of C. savignyi, which are usually found around plants that characterize typical dry steppe areas. As the ants' foraging paths recorded in the sympatric area reveal, C. bicolor performs significantly shorter foraging runs with respect to both length and time, and covers a much smaller foraging range than C. savignyi does. This result reflects the fact that the microhabitat occupied by the colonies of C. bicolor is richer in food abundance. When direct interspecific interactions were investigated by placing a bait midway between two heterospecific nests, C. bicolor foragers dominated over those of C. savignyi. The same dominance of C. bicolor over C. savignyi occurred in laboratory experiments. These results suggest that the dominant species drives the subordinate one out of the high quality microhabitats, and that the subordinate species is forced to survive in the less lucrative habitats. In conclusion, coexistence seems to be maintained by the asymmetric competitive relationship between the two species and the fact that the subordinate species has the ability to endure in the less favourable microhabitat.

Animals↗

A field test for competitive effects of Aedes albopictus on A. aegypti in South Florida: differences between sites of coexistence and exclusion?

We tested whether interspecific competition from Aedes albopictus had measurable effects on A. aegypti at the typical numbers of larval mosquitoes found in cemetery vases in south Florida. We also tested whether the effect of interspecific competition from A. albopictus on A. aegypti differed between sites where A. aegypti either persists or went extinct following invasion by A. albopictus. Similar experiments manipulating numbers of A. albopictus in cemetery vases were conducted at three sites of A. aegypti persistence and three sites where A. aegypti was apparently extinct. The experiments were done using numbers of larvae that were determined by observed numbers of larvae for each site, and with resources (leaf detritus) that accumulated in experimental vases placed into each field site. In both the early rainy season (when number of mosquito larvae was low) and the late rainy season (when number of mosquito larvae was high), there was a significant effect of treatment on developmental progress of experimental A. aegypti. In the late rainy season, when numbers of larvae were high, there was also a significant effect of treatment on survivorship of A. aegypti. However, the competition treatment x site type (A. aegyptipersists vs extinct) interaction was never significant, indicating that the competitive effect of A. albopictus on A. aegypti did not differ systematically between persistence versus extinction sites. Thus, although competition from A. albopictus is strong under field conditions at all sites, we find no evidence that variation in the impact of interspecific competition is associated with coexistence or exclusion. Interspecific competition among larvae is thus a viable explanation for exclusion or reduction of A. aegypti in south Florida, but variation in the persistence of A. aegypti following invasion does not seem to be primarily a product of variation in the conditions in the aquatic environments of cemetery vases.

Aedes↗

Competitive interactions in macroinfaunal animals of exposed sandy beaches.

The influence of biotic interactions in structuring macroinfaunal communities of exposed sandy beaches, an unstable habitat characterized by strong physical forces, is generally considered negligible. We investigated the hypothesis that competitive interactions during burrowing could potentially affect the intertidal distribution and abundance of macroinfaunal animals of sandy beaches using two species of invertebrates, a hippid crab, Emerita analoga, and a bivalve, Mesodesma donacium, common along the coast of Chile. Spatial overlap in the intertidal distributions of these species was dynamic, varying with abundance, location, time of year and tide. Highest density zones of each species were often distinctly separated at low tide and spatial overlap in their distributions decreased significantly with increasing density, suggesting density dependence of the interactions. Negative relationships between densities of the two species at the smallest spatial scale examined also suggested active interactions among individuals. Over a tidal cycle, peak densities of the two species overlapped suggesting that interactions could occur frequently. Burrowing performance of E. analoga varied between size classes in three experimental densities of clams (5, 10 and 15 clams 0.008 m(-2)) and in controls with no clams. Burrowing times of large crabs were significantly longer (approximately twofold) in all densities of clams than in controls, while those of small crabs did not differ significantly among treatments and controls. Large crabs also displaced clams from the sand while burrowing suggesting that two mechanisms of direct interference can occur, both of which could increase exposure of individuals involved to active swash and transport across or along the beach with potentially negative consequences. Our results suggest that competitive interactions capable of affecting zonation and population and community biology on a number of scales can occur among burrowing macroinfauna on exposed sandy beaches. Those interactions could be more ecologically significant than previously appreciated and may contribute to patterns observed in community structure and zonation on sandy beaches. Our results illustrate the potential importance of negative biological interactions in a physically stressful environment.

Analysis of Variance↗

Intraguild predation and interference competition on the endangered dragonfly Aeshna viridis.

We examined the effects of intraguild predation (IGP) and interference competition on an endangered dragonfly, Aeshna viridis Eversm. (Odonata: Anisoptera). A. viridis is rare in Europe due to the decrease in suitable habitats harboring the macrophyte Stratiotes aloides L. Stratiotes plants are the principal oviposition substrate for A. viridis females and protect the larvae of A. viridis from fish predation. In our study lakes A. viridis larvae are sympatric with larvae of Aeshna grandis and Aeshna juncea. The susceptibility of A. viridis larvae to IGP by similar-sized larvae of A. grandis and A. juncea was tested in a laboratory predation experiment. Microhabitat use of A. viridis and A. grandis was studied in the laboratory to determine the possible effects of interference competition on the spatial distribution of A. viridis larvae. Our results show that at least in laboratory conditions, A. viridis is susceptible to IGP and interference competition. In competition, A. grandis larvae dominated the middle and outer portion of S. aloides rosettes whereas A. viridis stayed in the inner parts. When A. grandis larvae were absent, A. viridis colonized the middle and outer parts of the rosettes. We conclude that asymmetric predation between odonate larvae of equal size can be intense, and that both IGP and interference competition affect A. viridis. Although natural habitat complexity diminishes their impact, these interactions may nevertheless influence the distribution of A. viridis in S. aloides waters and restrict its microhabitat use in S. aloides rosettes.

Animals↗

Influence of recruit condition on food competition and predation risk in a coral reef fish.

Settlement rate is considered to be a major determinant of the population structure of coral reef fishes. In this study, the effects of larval physiological condition on survival, predation risk and competitive ability are assessed for a small damselfish, Pomacentrus moluccensis. New settlers were collected and fed for 5 days to produce high and low condition (measured as lipid) treatment fish. In a field experiment, pairs (one high and one low condition fish) were transplanted to corals. Persistence over 2 weeks was much higher (100% vs. 25%) in high condition fish. In mixed groups in the laboratory, high condition fish were both aggressively dominant and consumed more of a limiting prey source than low condition fish. In addition, low condition fish were shown to be at much higher risk of predation. All of the low condition fish but only 33% of high condition fish in mixed groups were consumed by fish predators, and in a separate experiment, 73% of feeding strikes by predators were directed at low condition fish. Quality of new settlers can have an important influence on subsequent juvenile survival. The mechanisms for this effect are likely to include a combination of effects of condition on food competition and predation risk.

Animals↗

Intraguild predation and successful invasion by introduced ladybird beetles.

Introductions of two ladybird beetle (Coleoptera: Coccinellidae) species, Coccinella septempunctata and Harmonia axyridis, into North America for aphid biocontrol have been followed by declines in native species. We examined intraguild predation (IGP) between larvae of these two exotic species and larvae of the two most abundant native coccinellids in eastern Washington State, C. transversoguttata and Hippodamia convergens. In pairings between the two native species in laboratory microcosms containing pea ( Pisum sativum) plants, neither native had a clear advantage over the other in IGP. When the natives were paired with either Harmonia axyridis or C. septempunctata, the natives were more frequently the victims than perpetrators of IGP. In contrast, in pairings between the exotic species, neither had an IGP advantage, although overall rates of IGP between these two species were very high. Adding alternative prey (aphids) to microcosms did not alter the frequency and patterns of relative IGP among the coccinellid species. In observations of encounters between larvae, the introduced H. axyridis frequently survived multiple encounters with the native C. transversoguttata, whereas the native rarely survived a single encounter with H. axyridis. Our results suggest that larvae of the native species face increased IGP following invasion by C. septempunctata and H. axyridis, which may be contributing to the speed with which these exotic ladybird beetles displace the natives following invasion.

Animals↗

Differential effects of habitat complexity, predators and competitors on abundance of juvenile and adult coral reef fishes.

Greater structural complexity is often associated with greater abundance and diversity, perhaps because high complexity habitats reduce predation and competition. Using 16 spatially isolated live-coral reefs in the Bahamas, I examined how abundance of juvenile (recruit) and adult (non-recruit) fishes was affected by two factors: (1) structural habitat complexity and (2) the presence of predators and interference competitors. Manipulating the abundance of low and high complexity corals created two levels of habitat complexity, which was cross-factored with the presence or absence of resident predators (sea basses and moray eels) plus interference competitors (territorial damselfishes). Over 60 days, predators and competitors greatly reduced recruit abundance regardless of habitat complexity, but did not affect adult abundance. In contrast, increased habitat complexity had a strong positive effect on adult abundance and a weak positive effect on recruit abundance. Differential responses of recruits and adults may be related to the differential effects of habitat complexity on their primary predators. Sedentary recruits are likely most preyed upon by small resident predators that ambush prey, while larger adult fishes that forage widely and use reefs primarily for shelter are likely most preyed upon by large transient predators that chase prey. Increased habitat complexity may have inhibited foraging by transient predators but not resident predators. Results demonstrate the importance of habitat complexity to community dynamics, which is of concern given the accelerated degradation of habitats worldwide.

Analysis of Variance↗

Complex interactions between native and invasive fish: the simultaneous effects of multiple negative interactions.

We suggest that the ultimate outcome of interactions between native species and invasive species (extinction or coexistence) depends on the number of simultaneous negative interactions (competition and predation), which depends on relative body sizes of the species. Multiple simultaneous interactions may constrain the ability of native species to trade fitness components (i.e., reduced growth for reduced risk of predation) causing a spiral to extinction. We found evidence for five types of interactions between the adults and juveniles of introduced western mosquitofish (Gambusia affinis) and the juveniles of native least chub (Iotichthys phlegethontis). We added ten large (23-28 mm) and seven small (9-13 mm) young-of-the-year (YOY) least chub to replicate enclosures with zero, low, and high densities of mosquitofish in a desert spring ecosystem. Treatments with mosquitofish reduced the average survival of least chub by one-third. No small YOY least chub survived in enclosures with high mosquitofish densities. We also performed two laboratory experiments to determine mortality to predation, aggressiveness, and habitat selection of least chub in the presence of mosquitofish. Mean mortality of least chub due to predation by large mosquitofish was 69.7% over a 3-h trial. Least chub were less aggressive, selected protected habitats (Potamogeton spp.), and were more stationary in the presence of mosquitofish where the dominance hierarchy was large mosquitofish>>large least chub approximately small mosquitofish>>small least chub. Least chub juveniles appear to be figuratively caught in a vice. Rapid growth to a size refuge could reduce the risk of predation, but the simultaneous effects of competition decreased least chub growth and prolonged the period when juveniles were vulnerable to mosquitofish predation.

Analysis of Variance↗

Competitive equivalence maintains persistent inter-clonal boundaries.

Clear boundaries often separate adjacent conspecific competitors. These boundaries may reflect bordering animal territories or regions of inter-organism contact in mobile and non-mobile organisms, respectively. Sessile, clonal organisms often form persistent inter-clonal boundaries despite great variation in competitive ability among genotypes within a population. I show that neighboring clones in the sea anemone Anthopleura elegantissima and three species of the marine hydroid genus Hydractinia are more evenly matched in terms of competitive ability than expected by chance. Hypotheses of genetic relatedness or similar environmental regime shared by neighboring clones are inconsistent with the observed similarities between adjacent competitors in one or both taxa. Instead, inter-clonal borders evidently persist as standoffs between evenly matched competitors. Large differences in competitive ability between bordering clones were rarely observed, suggesting that dominant clones quickly displace or eliminate others in competitive mismatches. This ecological parallel between taxa (i.e., competitive equivalence) exists despite several fundamental differences (e.g., geographical distribution, habitat, body size, longevity), suggesting that competitive equivalence may be a widespread determinant of boundary persistence between adjacent competitors.

Analysis of Variance↗

Dynamics of demographically open mutualists: immigration, intraspecific competition, and predation impact goby populations.

Although it is now recognized that mutualistic species are common and can have stable populations, the forces controlling their persistence are poorly understood. To better understand the mechanisms that impact the stability of obligate mutualists, I conducted several field experiments within a sandy coral reef lagoon in Moorea, French Polynesia that manipulated densities of fish (gobies) that interact mutualistically with shrimp. Obligate, mutualistic partnerships of gobies and shrimp are common on Indo-Pacific coral reefs and have been shown previously to interact as follows: shrimp construct burrows in which both species reside, and gobies warn shrimp of predators through tactile communication. Augmentation of gobies by up to 100% above ambient densities within 9 m2 plots produced no change in overall density of gobies or shrimp because gobies competed intraspecifically for a limited number of shrimp burrows and smaller gobies were outcompeted by larger individuals. I used predators to assess the impact of goby removal on the stability of goby and shrimp populations. First, although surveys taken throughout the lagoon revealed no relationship between goby and predator densities, predators correlated negatively with the proportion of adult gobies and positively with the proportion of small gobies paired with large shrimp. Second, experimental augmentation of predators resulted in a dramatic reduction of adult gobies within predator-addition plots, but had no impact on overall densities as immigrants rapidly replaced the missing adult gobies. Furthermore, goby turnover resulted in an increase in the proportion of small gobies paired with large shrimp because body sizes of gobies and shrimp in a burrow were similar prior to predator introduction, and predators apparently had a greater impact on gobies than shrimp. The mechanisms that prevent expansion (intraspecific competition) and collapse (immigration) of goby-shrimp populations likely contribute to local-scale stability of mutualistic populations in other terrestrial and aquatic environments.

Animals↗

Experimental small-scale grassland fragmentation alters competitive interactions among ant species.

Different species may respond differently to habitat fragmentation. Theory predicts that abundant generalist species should be less affected by fragmentation than specialist species. In ant communities, the most abundant species is often behaviourally dominant. Thus, habitat fragmentation could alter competitive interactions between the dominant ant species and the other species. We tested this hypothesis in a long-term grassland fragmentation experiment. Fragments of different size (20.25 and 2.25 m(2)) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots were situated in adjacent undisturbed grassland. Ant density and species composition were assessed 3 and 6 years after initiation of the experimental fragmentation. The effect of the dominant ant species on the resource use of the other species was examined at natural sugar resources (aphids and extrafloral nectaries) and at artificial sugar baits. Lasius paralienus was the most abundant ant species (72% of nests) in the grasslands examined. Species richness and forager density in the other species decreased with increasing density of L. paralienus in fragments but not in control plots. The overall forager density of the other species was positively related to their habitat niche overlap with L. paralienus. The density of foragers of the other species at sugar resources was not affected by L. paralienus forager density. The experimental fragmentation resulted in an increase in natural sugar resources in fragments. This may have reduced the intensity of interspecific competition for sugar resources. Our study shows that the grassland fragmentation altered interactions between the dominant L. paralienus and the other ant species.

Analysis of Variance↗

Sheep grazing and rodent populations: evidence of negative interactions from a landscape scale experiment.

Inter-specific competition, facilitation and predation influence herbivore assemblages, but no study has experimentally explored the interactions between large ungulates and small rodents. In a fully replicated, landscape scale experiment, we manipulated densities of domestic sheep in mountain pastures in Norway. We then determined population growth and densities of rodents by live trapping in each of the areas with different sheep densities. We found that the (summer) population growth rate and autumn density of the field vole (Microtus agrestis) was lower at high sheep density. This provides the first experimental evidence of negative interactions between an ungulate and small rodent species. There was no effect on the bank vole (Clethrionomys glareolus), whose diet differs from sheep. Sheep density, therefore, potentially alters the pattern of inter-specific population synchrony amongst voles. Our study shows that negative interactions between large ungulates and small rodents may be species-specific and negative population consequences for the rodent population appear above threshold ungulate densities.

Animals↗

Do multitrophic interactions override N fertilization effects on Operophtera larvae?

We examined how performance of Operophtera brumata (Lepidoptera) larvae was affected by nitrogen (N) fertilization of boreal forest understorey vegetation. We monitored larval densities on Vaccinium myrtillus plants for a period of 7 years in a field experiment. Preliminary results indicated that the N effect on larval densities was weak. To examine if this was due to indirect interactions with a plant pathogen, Valdensia heterodoxa, that share the same host plant, or due to top-down effects of predation, we performed both a laboratory feeding experiment (individual level) and a bird exclusion experiment (population level) in the field. At the individual level, altered food plant quality (changes in plant concentration of carbon, N, phenolics, or condensed tannins) due to repeated infection by the pathogen had no effect on larval performance, but both survival to the adult stage and adult weight were positively affected by N fertilization. Exclusion of insectivorous birds increased the frequency of larval damage on V. myrtillus shoots, indicating higher larval densities. This effect was stronger in fertilized than in unfertilized plots, indicating higher bird predation in fertilized plots. Predation may thus explain the lack of fertilization effect on larval densities in the field experiment. Our results suggest that top-down effects are more important for larval densities than bottom-up effects, and that bird predation may play an important role in population regulation of O. brumata in boreal forests.

Analysis of Variance↗

Genotype x environment interactions, stoichiometric food quality effects, and clonal coexistence in Daphnia pulex.

The role of stoichiometric food quality in influencing genotype coexistence and competitive interactions between clones of the freshwater microcrustacean, Daphnia pulex, was examined in controlled laboratory microcosm experiments. Two genetically distinct clones of D. pulex, which show variation in their ribosomal rDNA structure, as well as differences in a number of previously characterized growth-rate-related features (i.e., life-history features), were allowed to compete in two different arenas: (1) batch cultures differing in algal food quality (i.e., high vs. low carbon:phosphorus (C:P ratio) in the green alga, Scenedesmus acutus); (2) continuous flow microcosms receiving different light levels (i.e., photosynthetically active radiation) that affected algal C:P ratios. In experiment 1, a clear genotype x environment interaction was determined with clone 1 out-competing clone 2 under high nutrient (i.e., low food C:P) conditions, while the exact opposite pattern was observed under low nutrient (i.e., high C:P) conditions. In experiment 2, clone 1 dominated over clone 2 under high light (higher C:P) conditions, but clonal coexistence was observed under low light (low C:P) conditions. These results indicate that food (nutrient) quality effects (hitherto an often overlooked factor) may play a role in microevolutionary (genotypic) responses to changing stoichiometric conditions in natural populations.

Analysis of Variance↗

Outcomes of reciprocal invasions between genetically diverse and genetically uniform populations of Daphnia obtusa (Kurz).

Ecological theory predicts that genetic variation produced by sexual reproduction results in niche diversification and provides a competitive advantage both to facilitate invasion into genetically uniform asexual populations and to withstand invasion by asexual competitors. We tested the hypothesis that a large group of diverse clones of Daphnia obtusa has greater competitive advantage when invading into genetically uniform populations of this species than a smaller group with inherently less genetic diversity. We compared competitive outcomes to those of genetically uniform groups of small and large size invading into genetically diverse populations. Genetically diverse invaders of initially large group size increased their representation by more than those of initially small size; in contrast, genetically uniform invaders of initially large group size diminished on average by more than those of initially small size. These results demonstrate an advantage to the genetic variation produced by sexual reproduction, both in invasion and resisting invasion, which we attribute to competitive release experienced by individuals in genetically diverse populations.

Animals↗

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of Apeiba membranacea.

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.

Animals↗

Nest site attributes and temporal patterns of northern flicker nest loss: effects of predation and competition.

To date, most studies of nest site selection have failed to take into account more than one source of nest loss (or have combined all sources in one analysis) when examining nest site characteristics, leaving us with an incomplete understanding of the potential trade-offs that individuals may face when selecting a nest site. Our objectives were to determine whether northern flickers (Colaptes auratus) may experience a trade-off in nest site selection in response to mammalian nest predation and nest loss to a cavity nest competitor (European starling, Sturnus vulgaris). We also document within-season temporal patterns of these two sources of nest loss with the hypothesis that flickers may also be constrained in the timing of reproduction under both predatory and competitive influence. Mammalian predators frequently depredated flicker nests that were: lower to the ground, less concealed by vegetation around the cavity entrance and at the base of the nest tree, closer to coniferous forest edges and in forest clumps with a high percentage of conifer content. Proximity to coniferous edges or coniferous trees increased the probability of nest predation, but nests near conifers were less likely to be lost to starlings. Flickers may thus face a trade-off in nest site selection with respect to safety from predators or competitors. Models suggested that peaks of nest predation and nest loss to eviction occurred at the same time, although a competing model suggested that the peak of nest loss to starlings occurred 5 days earlier than the peak of mammalian predation. Differences in peaks of mammalian predation and loss to starlings may constrain any adjustment in clutch initiation date by flickers to avoid one source of nest loss.

Animals↗