The effects of competition on fitness depend on the sex of both competitors

In intraspecific competition, the sex of competing individuals is likely to be important in determining the outcome of competitive interactions and the way exposure to conspecifics during development influences adult fitness traits. Previous studies have explored differences between males and females in their response to intraspecific competition. However, few have tested how the sex of the competitors, or any interactions between focal and competitor sex, influences the nature and intensity of competition. We set up larval seed beetles Callosobruchus maculatus to develop either alone or in the presence of a male or female competitor and measured a suite of traits: development time, emergence weight; male ejaculate mass, copulation duration, and lifespan; and female lifetime fecundity, offspring egg–adult survival, and lifespan. We found effects of competition and competitor sex on the development time and emergence weight of both males and females, and also of an interaction between focal and competitor sex: Females emerged lighter when competing with another female, while males did not. There was little effect of larval competition on male and female adult fitness traits, with the exception of the effect of a female competitor on a focal female's offspring survival rate. Our results highlight the importance of directly measuring the effects of competition on fitness traits, rather than distant proxies for fitness, and suggest that competition with the sex with the greater resource requirements (here females) might play a role in driving trait evolution. We also found that male–male competition during development resulted in shorter copulation times than male–female competition, a result that remained when controlling for the weight of competitors. Although it is difficult to definitively tease apart the effects of social environment and access to resources, this result suggests that something about the sex of competitors other than their size is driving this pattern.     

Effects of environmental variation on fitness of singly and multiply sired progenies of Raphanus sativus (Brassicaceae)

Two hypotheses have been proposed concerning possible fitness advantages of multiple paternity. According to the Elbow Room hypothesis, the magnitude of resource partitioning is positively correlated with the genetic diversity of competitors. This leads to the prediction that the mean fitness of competing half-siblings will exceed the mean fitness of competing full-siblings. The Lottery hypothesis suggests that in a patchy environment a genetically diverse sibship increases the probability that a mother will produce winning phenotypes for each of several different microsites. Both hypotheses were tested in a greenhouse experiment with wild radish, Raphanus sativus. Progeny derived from factorial crosses were raised in competition as full- or half-siblings. To simulate environmental variation, the experiment was replicated across a gradient of nutrient concentrations. After 9 weeks, the aboveground biomass of all plants was harvested and oven-dried. Although dry weights were strongly influenced by nutrient level, within nutrient classes there were no significant differences in the mean dry weight of full- and half-sib competitors. Thus, there was no evidence for the Elbow Room hypothesis. In addition, there were no significant differences in numbers of winning phenotypes in the two competition regimes. Therefore, there was also no support for the Lottery hypothesis. Although competition regime did not influence the mean fitness of competing progeny, it significantly affected variation in dry weight of seedlings within pots. Higher coefficients of variation in half-sib pots suggest that interference competition may be greater in genetically diverse sibships.     

Details of local dispersion improve the fit of neighborhood competition models

I performed a series of greenhouse experiments to explore how patterns in the dispersion of local competitors affect the reproductive performance of Capsella bursa-pastoris, Poa annua and Senecio vulgaris. I manipulated the density and relative frequency of competitors in each of three concentric rings surrounding a central plant, thereby creating a variety patterns by which local competitors were distributed. Neighborhood competition models were used to predict the seed output of these central plants. For both Senecio vulgaris and Capsella bursa-pastoris, I found that models which incorporated the dispersion of competitors, as well as the relative emergence date of plants, performed substantially better than those that considered only the distance between the central plant and each of its competitors. I was unable to measure the seed output of Poa annua but neither emergence data nor the dispersion of competitors were important in determining its final dry-weight. Present address: Department of Biology, Box 1137, Washington University, 1 Brookings Drive, St. Louis, MO 63130, USA     

Competition along a spatial gradient of resource supply: a microbial experimental model

In a set of laboratory experiments, we examined competition for phosphorus between algae and bacteria under various carbon:phosphorus (C:P) supply ratios in spatially homogeneous and heterogeneous microcosms. Experimental results were compared to those predicted by theoretical models of resource competition. In the spatially heterogeneous microcosm, algae that were inferior competitors for P persisted in vessels with high local C:P supply ratios that would cause exclusion in the spatially homogeneous microcosms. Resource competition theory, adapted to this system, provided a starting point for explaining these results. Spatial structure can enhance local diversity because locally inferior competitors are transported from source habitats into sink habitats where they would otherwise be excluded. Such local sources were determined by their resource supply ratios. These results verify the hypothesis that spatial processes enhance local diversity when a system of local habitats is divided into sources and sinks in such a way that each persisting species has at least one source within the system. However, existing theoretical models did not accurately predict distributions of competitor abundance within this experimental system.     

FITNESS CONSEQUENCES OF MATING SYSTEM,SEED WEIGHT,AND EMERGENCE DATE IN A WINTER ANNUAL,COLLINSIA VERNA

Selfed and outcrossed progeny of 60 maternal parents were produced to investigate the joint and individual effects of mating system, seed weight, and emergence date on the expression of characters related to fitness and adult fecundity. A series of analyses of variance investigated these effects through time and indicated that 1) mating system explained 56% of the variance in seed weight, 2) seed weight explained 51% and mating system explained 38% of the variance in emergence date, and 3) mating system explained 71% and seed weight explained 15% of the variance in fecundity. Outcrossed-seed means differed significantly from selfed-seed means for all traits measured. On average, outcrossed seeds were larger, germinated earlier, had higher percentage emergence, and produced plants that were more fecund than selfed seeds. The coefficient of inbreeding depression increased through time in this study, from 0.05 for seed weight to 0.23 for fecundity. Seed weight and emergence date were positively correlated, both phenotypically and genetically, for both mating systems. Genetically, this indicates that genes that increase the value of seed weight also increase the value of emergence date and vice versa. Phenotypically, the positive correlation indicates that larger seeds germinate later. Outcrossed seeds were significantly larger but germinated earlier than selfed seeds, suggesting that mating system has an overriding effect in influencing fitness. In light of the selection on emergence date quantified in a previous study, seed weight, emergence date, and mating system may be functioning as a cluster of characters on which selection acts jointly in this species.     

Differences in life history strategies between an invasive and a competing resident seed predator

Competitive interactions may arise from biological invasion if a successful invasive species requires the ecological niche of a resident one. Life-history traits that make a species a successful invader are of particular interest in elucidating both invasion success and how interspecific competition may emerge. In southeastern France, the invasion of cedar forests by the seed chalcid Megastigmus schimitscheki generated competitive relationships with the resident M. pinsapinis for the exploitation of the seed resource. Among the numerous ecological traits allowing these seed predators to exploit their niche successfully, the timing of adult emergence, initial egg load and age-specific realized fecundities of females were investigated to help understanding the issue of such interspecific relationships. Spring adult emergence of M. schimitscheki under natural conditions was significantly earlier than that of M. pinsapinis, suggesting that an advantage for the access to the seed resource for the invasive species may be associated with this trait. Initial egg load was significantly higher in M. schimitscheki than in M. pinsapinis and the analysis of age-specific realized fecundity in semi-natural conditions indicated that both M. schimitscheki and M. pinsapinis females lay a large proportion of their eggs during the early days of their lives. In the light of these findings, both earlier phenology and higher reproduction abilities of M. schimitscheki may have the potential to confer an advantage within a competitive context with M. pinsapinis through enhanced seed resource preemption. This may potentially explain the invasiveness of M. schimitscheki in southern France despite the presence of its closely related competitor M. pinsapinis.     

Asymmetric Competition Among Distant Taxa

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

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.     

Female philopatry and male-biased dispersal in a direct-developing salamander, Plethodon cinereus

The local resource competition hypothesis and the local mate competition hypothesis were developed based on avian and mammalian systems to explain sex-biased dispersal. Most avian species show a female bias in dispersal, ostensibly due to resource defence, and most mammals show a male bias, ostensibly due to male-male competition. These findings confound phylogeny with mating strategy; little is known about sex-biased dispersal in other taxa. Resource defence and male-male competition are both intense in Plethodon cinereus, a direct-developing salamander, so we tested whether sex-biased dispersal in this amphibian is consistent with the local resource competition hypothesis (female-biased) or the local mate competition hypothesis (male-biased). Using fine-scale genetic spatial autocorrelation analyses, we found that females were philopatric, showing significant positive genetic structure in the shortest distance classes, with stronger patterns apparent when only territorial females were tested. Males showed no spatial genetic structure over the shortest distances. Mark-recapture observations of P. cinereus over 5 years were consistent with the genetic data: males dispersed farther than females during natal dispersal and 44% of females were recaptured within 1 m of their juvenile locations. We conclude that, in this population of a direct-developing amphibian, females are philopatric and dispersal is male-biased, consistent with the local mate competition hypothesis.     

Effects of spatial pattern and relatedness in an experimental plant community

Many plant species show limited dispersal resulting in spatial and genetic substructures within populations. Consequently, neighbours are often related between each other, resulting in sibling competition. Using seed families of the annuals Capsella bursa-pastoris and Stachys annua we investigated effects of spatial pattern (i.e. random versus aggregated) on total and individual performance at the level of species and seed families under field conditions. At the level of species, we expected that inferior competitors increase, while superior competitors decrease their performance within neighbourhoods of conspecifics. Thus, we expected a species by spatial pattern interaction. Sibling competition, however, might reduce the performance of competitors, when genetically related, rather than non-related individuals are competing. Therefore, aggregations at the level of seed families could decrease the performance of competitors. Alternatively, if the opposite outcome would be observed, kin selection might be hypothesized to have occurred in the past. Because heavy seeds are expected to disperse less than light seeds, we further hypothesized that kin selection might be more likely to occur in superior competitors with heavy, locally dispersed seeds (e.g. Stachys) compared to inferior competitors with light, more distantly dispersed seeds (e.g. Capsella). We found a significant species by spatial pattern interaction. Indeed, the inferior competitor, Capsella, showed increased reproductive biomass production in aggregated compared to random patterns. Whereas, the performance of the superior competitor, Stachys, was to some extent decreased by intraspecific aggregation. Although statistically not significant, effects of intrafamily aggregations tended to be rather negative in Capsella but positive in Stachys. Our results confirmed that spatial patterns affect growth and reproduction of plant species promoting coexistence in plant communities. Although, we could not provide strong evidence for sibling competition or kin selection, our results suggested that competition among relatives was more severe for Capsella (lighter seeds) compared to Stachys (heavier seeds).     

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

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

ENHANCEMENT OF INBREEDING DEPRESSION BY DOMINANCE AND SUPPRESSION IN IMPATIENS CAPENSIS

We investigated the effect of intraspecific competition on the magnitude of inbreeding depression in Impatiens capensis by planting seeds from chasmogamous (CH) and cleistogamous (CL) flowers in three experimental greenhouse treatments: in individual pots, in flats in dense pure stands according to seed type, and in flats with the two seed types intermixed in a checkerboard array. The size distributions of plants grown in flats were significantly more hierarchical than those of plants grown individually, indicating that larger plants competitively suppressed smaller plants in the high-density treatments. The magnitude of inbreeding depression at high density depended upon the planting arrangement of CL and CH seeds. CH advantage was greatest when CH and CL seedlings were grown in competition with one another, suggesting that fitness differences between outcrossed and inbred individuals were intensified by dominance and suppression. For plants grown individually, the effects of maternal parent, seed weight, and emergence date on seedling size disappeared with plant age, whereas at high density these effects remained at the final harvest. Thus, plant density may influence patterns of natural selection both on mating system and on juvenile traits in natural Impatiens populations.     

Odour-mediated long-range avoidance of interspecific competition by a solitary endoparasitoid: a time-saving foraging strategy

1. In studies on optimal foraging strategies, long-range decisions in the pursuit of resource are rarely considered. This is also the case for sympatric parasitoids, which may be confronted with the decision to accept or reject host larvae that are already parasitized by a competing species. They can be expected to reject already parasitized hosts if it is likely that they will lose the resulting intrinsic competition. However, examples of such interspecific host discrimination are rare. 2. We propose that parasitoids that are not egg-limited should reject inferior hosts only if it saves them time, and that this will be achieved mainly when the parasitoids are able to detect competitors from a distance. We tested this hypothesis using the sympatric parasitoids Cotesia marginiventris (Cresson) and Campoletis sonorensis (Cameron). 3. C. sonorensis was found to be the superior intrinsic competitor but, upon contact with a host larva, both wasps readily accepted hosts that had already been parasitized by the other species. However, in an olfactometer experiment, C. marginiventris females were found to strongly avoid the odour of their superior competitor. 4. These results are in accordance with a time optimization scenario, whereby the inferior competitor accepts competition if it costs only an egg, but avoids competition if it may save time that can be allocated to the search for more profitable hosts. 5. Models on host discrimination strategies in parasitoids had not yet considered discrimination from a distance. Long-range foraging decisions can also be expected for other organisms that have to choose between resources of varying suitability and profitability.     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

Gene differences in heading date,height, seed weight and seed yield between two pure line varieties of Triticum aestivum L.

Summary Reciprocal sets of homozygous inbred backcross lines were developed by crossing two pure line varieties (Baart 46 and Ramona) of Triticum aestivum L., followed by two backcrosses to each of the two parent varieties, and six to eight generations of selfing. Data on each inbred backcross line was obtained from twelve plots (from replications in three years). Five genes were responsible for over 95% of the genetic variation for heading date. These genes had pleiotropic effects on plant height that were proportional to their effects on heading date. Two additional genes had detectable effects on plant height. The genes with a measurable effect on height accounted for 90% of the genetic variation in the Baart 46 genetic background. One gene affected seed weight. In the Ramona background, this gene accounted for 80% of the genetic variation in seed weight and 16% of the genetic variation in seed yield. Two genes, responsible for the earliest and latest heading date classes, had large pleiotropic effects on seed yield. They accounted for 60% of the genetic variation in yield. One gene, with no effect on heading date, caused a detectable reduction in yield of 23% in the Baart 46 inbred backcross lines. This gene had no apparent effect in the Ramona genetic background. Quantitative trait genes are sparsely distributed in the genome: fewer than one in four chromosome arms carries a gene with a detectable effect. Gene effects on quantitative traits are not small and similar. The distribution of 22 gene effects for heading date and height is slightly skewed to the right: as the magnitude of effect increases, the frequency of genes having the effect decreases.     

Effects of multiple competitors for pollination on bumblebee foraging patterns and Mimulus ringens reproductive success

When co‐occurring plant species overlap in flowering phenology they may compete for the service of shared pollinators. Competition for pollination may lower plant reproductive success by reducing the number of pollinator probes or by decreasing the quality of pollen transport to or from a focal species. Pair‐wise interactions between plants sharing pollinators have been well documented. However, relatively few studies have examined interactions for pollination among three or more plant species, and little is known about how the outcomes and mechanisms of competition for pollination may vary with competitor species composition. To better understand how the dynamics of competition for pollination may be influenced by changes in the number of competitors, we manipulated the presence of two competitors, Lythrum salicaria and Lobelia siphilitica, and quantified reproductive success for a third species, Mimulus ringens. Patterns of pollinator preference and interspecific transitions in mixed‐species arrays were significantly influenced by the species composition of competitor plants present. Both pair‐wise and three‐species competition treatments led to a similar ～ 40% reduction in Mimulus ringens seed set. However, the patterns of pollinator foraging we observed suggest that the relative importance of different mechanisms of competition for pollination may vary with the identity and number of competitors present. This variation in mechanisms of competition for pollination may be especially important in diverse plant communities where many species interact through shared pollinators.     

EFFECT OF MALE POLLEN DONOR AND FEMALE SEED PARENT ON ALLOCATION OF RESOURCES TO DEVELOPING SEEDS AND FRUIT IN CHAMAECRISTA FASCICULATA (LEGUMINOSAE)

In Chamaecrista fasciculata, fruit abortion levels are high and seed mass is highly variable, necessary preconditions for differential resource allocation of the female to seed and fruit sired by different males. This study investigated the relative role of pollen donor and seed parent on the allocation of resouces to developing seed and fruit, and assessed the role of genetic relatedness in contributing to any observed paternal effect in C. fasciculata. In addition, pollen donor effects were contrasted to within-seed parent sources of variation in resource allocation due to pollination date and ovule position in the pod. Plants collected from the field were brought to a greenhouse where single-donor crosses were conducted controlling for pollen donor source and interplant distance, a measure of genetic relatedness. Seed mass, number of seed/fruit, fruit maturation time, and fruit abortion rate were measured as indicators of resource allocation to developing seed and fruit. Variation in resource allocation was largely determined by the seed parent. Pollen donor effects were limited to differences between self vs. non-self pollinations, suggesting that inbreeding depression following mating events between related individuals is the source of any variation among pollen donors on differential resource allocation to developing seed and fruit. Once the effect of inbreeding was removed, however, pollination date and ovule position played larger roles than pollen source. Since there was no detectable variation among male pollen donors in their ability to accrue resources from the female seed parent apart from inbreeding effects, it is concluded that the opportunity for postzygotic mate choice is limited in C. fasciculata.     

Competition and genetic background in a rapid-cycling cultivar of brassica rapa (Brassicaceae)

We examined the effects of a slight genetic disadvantage in a competitive situation by comparing the performance of inbred and outcrossed Brassica rapa plants over a range of neighbor densities, using a rapid-cycling, self-compatible cultivar of this species. We also examined the genetic correlations in performance among plants grown alone and with intra- and interspecific competitors. Competition had a strong effect on biomass and on the number of flowers produced, but differences in biomass between inbred and outcrossed plants were dependent on the competitive environment. Outcrossed plants outperformed those that were selfed only at intermediate densities of neighbors; at high densities and in the absence of competition inbred and outcrossed plants did not differ. For outcrossed families, performance without competitors and in intra- and interspecific competition were all highly correlated, but for selfed families, correlations were low, and performance in competition was not predictable from that of plants grown alone. Thus, the phenotypic expression of genetic differences may depend on the density of neighbors with which plants are grown.     

Maternal provisioning is structured by species’ competitive neighborhoods

Differential maternal provisioning of offspring in response to environmental conditions has been argued as ‘the missing link’ in plant life histories. Although empirical evidence suggests that maternal provisioning responses to abiotic conditions are common, there is little understanding of how differences in maternal provisioning manifest in response to competition. Frequency manipulations are commonly employed in ecological studies to assess the strength of interspecific competition, relative to intraspecific competition, and we used frequency manipulations to test how competition in two soil moisture environments affects maternal provisioning of seed mass. Specifically, for 15 pairs comprised from 25 annual plant species that occur in California, we varied the relative frequencies of conspecific to heterospecific competitors from 90% (intraspecific competition) to 10% (interspecific competition). We found that conspecific frequency affected maternal provisioning (seed mass) in 12 of the 25 species (eight significantly (p < 0.05), four marginally significantly (p < 0.07)), and that these responses included both increased (five species) and decreased (six species) seed mass, as well as one species with opposing directions of response to conspecific frequency that depended on the soil moisture environment. Conspecific frequency also affected per capita fecundity (seed number) for 17 of the 25 species (15 significantly (p < 0.05), two marginally significantly (p < 0.09)), which generally decreased seed number as conspecific frequency increased. The direction and magnitude of frequency‐dependent seed mass depended on the identity of the competitor, even among species whose fecundity was not affected by competitor identity; the latter finding reveals competitive differences among species that would otherwise appear to be competitively equivalent. Our research demonstrates how species responses to different competitive environments manifest through maternal provisioning, and that these responses alter previous estimates of environmentally‐determined maternal provisioning and reproductive output; future study is needed to understand their combined effects on population and community dynamics.