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PubMed2000

Mesoscale vortices and the paradox of the plankton.

Coexistence of competitive species is severely limited by the availability of resources and the characteristics of the environment. In particular, the so-called 'competitive exclusion principle' states that, at equilibrium, the number of coexisting species cannot be larger than the number of resources for which they compete. However, many in situ observations have revealed prolonged coexistence of a large number of competitive plankton species, a phenomenon known as 'the paradox of the plankton'. Here we investigate this problem and show that ocean mesoscale vortices generate transport barriers and incomplete horizontal mixing, allowing for a prolonged survival of the less-fit species, even for fully homogeneous resource distributions. In such a situation, the temporarily less-fit plankton species are protected from competition by the action of the vortices.

Animals↗

PubMed2000

Male-male competition and parental care in collared flycatchers (Ficedula albicollis): an experiment controlling for differences in territory quality.

Females are known to benefit from mate choice in several different ways but the relationship between these benefits has received little attention. The quality of resources provided by males, such as nest sites, and paternal care are often assumed to covary positively However, because the location of the nest affects the cost of parental care, these two benefits from mate choice can easily be confounded. To investigate the provisioning ability of successful competitors while controlling for differences in territory quality we removed early-settled pairs of collared flycatchers (Ficedula albicollis) and allowed replacement by later-arriving males or floaters (i.e.'poor competitors'). A control group of early-settled males (i.e. 'good competitors') had their females removed. Females paired to good competitors enjoyed a significantly higher reproductive success and tended to receive more parental assistance from their mates compared with females mated to poor competitors. Thus, some males seem able not only to compete successfully over resources but also to feed their offspring at a relatively higher rate. An alternative explanation, that poor competitors invested less in offspring quality in response to a lower share of paternity, could be rejected. The rate of extra-pair paternity did not differ between the two treatment groups. Our results suggest that male- male competition can sometimes facilitate female choice of superior care-givers. Thus, a female's benefit from choosing a competitive male may not be restricted to the quality of the resource he defends but can also include superior paternal care.

Animals↗

PubMed2001

Adjustment of offspring sex ratios in relation to the availability of resources for philopatric offspring in the common brushtail possum.

The local-resource-competition hypothesis predicts that where philopatric offspring compete for resources with their mothers, offspring sex ratios will be biased in favour of the dispersing sex. This should produce variation in sex ratios between populations in relation to differences in the availability of resources for philopatric offspring. However, previous tests of local resource competition in mammals have used indirect measures of resource availability and have focused on sex-ratio variation between species or individuals rather than between local populations. Here, we show that the availability of den sites predicts the offspring sex ratio in populations of the common brushtail possum. Female possums defend access to dens, and daughters, but not sons, occupy dens within their mother's range. However, the abundances of possums in our study areas were determined principally by food availability. Consequently, in food-rich areas with a high population density, the per-capita availability of dens was low, and the cost of having a daughter should have been high. This cost was positively correlated with male bias in the sex ratio at birth. Low per capita availability of dens was correlated with male bias in the sex ratio at birth.

Animals↗

PubMed2001

Mate guarding in the Seychelles warbler is energetically costly and adjusted to paternity risk.

Males may increase their fitness through extra-pair copulations (copulations outside the pair bond) that result in extra-pair fertilizations, but also risk lost paternity when they leave their own mate unguarded. The fitness costs of cuckoldry for Seychelles warblers (Acrocephalus sechellensis) are considerable because warblers have a single-egg clutch and, given the short breeding season, no time for a successful replacement clutch. Neighbouring males are the primary threat to a male's genetic paternity. Males minimize their loss of paternity by guarding their mates to prevent them from having extra-pair copulations during their fertile period. Here, I provide experimental evidence that mate-guarding behaviour is energetically costly and that the expression of this trade-off is adjusted to paternity risk (local male density). Free-living males that were induced to reduce mate guarding spent significantly more time foraging and gained significantly better body condition than control males. The larger the reduction in mate guarding, the more pronounced was the increase in foraging and body condition (accounting for food availability). An experimental increase in paternity risk resulted in an increase in mate-guarding intensity and a decrease in foraging and body condition, and vice versa. This is examined using both cross-sectional and longitudinal data. This study on the Seychelles warbler offers experimental evidence that mate guarding is energetically costly and adjusted to paternity risk.

Animals↗

PubMed2001

Skewed paternity and sex allocation in hermaphroditic plants and animals.

Models predict a reduced allocation to sperm when females preferentially use one of two males' sperm and the males do not know who is favoured. An analogous discounting occurs in plants when their paternity success is skewed by random, non-heritable factors such as location in the population and pollinator behaviour. We present a model that shows that skewed paternity can affect the sex allocation of hermaphrodites, that is it leads to a female-biased investment. The model highlights the close links between local mate competition and sperm competition. We use paternity data from Ficus in order to illustrate that skews in paternity success can lead to a high degree of sibling gamete competition in an apparently open breeding system. Since skews in paternity are ubiquitous in hermaphroditic plants and animals these findings should apply broadly.

Animals↗

PubMed2001

Evolutionary disarmament in interspecific competition.

Competitive asymmetry, which is the advantage of having a larger body or stronger weaponry than a contestant, drives spectacular evolutionary arms races in intraspecific competition. Similar asymmetries are well documented in interspecific competition, yet they seldom lead to exaggerated traits. Here we demonstrate that two species with substantially different size may undergo parallel coevolution towards a smaller size under the same ecological conditions where a single species would exhibit an evolutionary arms race. We show that disarmament occurs for a wide range of parameters in an ecologically explicit model of competition for a single shared resource; disarmament also occurs in a simple Lotka-Volterra competition model. A key property of both models is the interplay between evolutionary dynamics and population density. The mechanism does not rely on very specific features of the model. Thus, evolutionary disarmament may be widespread and may help to explain the lack of interspecific arms races.

Adaptation, Biological↗

PubMed2002

Invasion of vacant niches and subsequent sympatric speciation.

An individual-based simulation study was conducted to examine the population dynamics of 'invasion of a vacant niche' and subsequent speciation (by reproductive isolation) when food resources are randomly distributed spatially within the habitat and the frequencies of different food types are bimodally distributed (i.e. smaller and larger sizes of food being most abundant). The initially vacant niche was that of unused larger sizes of food. When phenotypic variation for resource use (i.e. food sizes) was small in the initial population, and each female could choose a mate from anywhere in the habitat, the population could not invade the vacant niche. But when the dispersal distance of the offspring and the area within which a female could choose a mate were small (i.e. the genetic neighbourhood size was small), the population could, in most cases, evolve to use both smaller and larger food sizes and form sister species sympatrically, with each species utilizing one of the two niches (small and large sizes of food). When phenotypic variation in resource use in the initial population was large, the population could, in most cases, invade the vacant niche by evolving to use both smaller and larger sizes of food. The probability of speciation increased as the dispersal distance of offspring decreased. The results indicate that populations whose individuals have small Wright's genetic neighbourhoods may often exploit a vacant niche and diversify sympatrically in the process.

Animals↗

PubMed2002

Chimpanzees and the mathematics of battle.

Recent experiments have demonstrated the importance of numerical assessment in animal contests. Nevertheless, few attempts have been made to model explicitly the relationship between the relative number of combatants on each side and the costs and benefits of entering a contest. One framework that may be especially suitable for making such explicit predictions is Lanchester's theory of combat, which has proved useful for understanding combat strategies in humans and several species of ants. We show, with data from a recent series of playback experiments, that a model derived from Lanchester's 'square law' predicts willingness to enter intergroup contests in wild chimpanzees (Pan troglodytes). Furthermore, the model predicts that, in contests with multiple individuals on each side, chimpanzees in this population should be willing to enter a contest only if they outnumber the opposing side by a factor of 1.5. We evaluate these results for intergroup encounters in chimpanzees and also discuss potential applications of Lanchester's square and linear laws for understanding combat strategies in other species.

Aggression↗

PubMed2002

Selection for restraint in competitive ability in spatial competition systems.

The absence of 'super competitors' in nature is usually attributed to organisms facing trade-offs in resource allocation. Here we identify another mechanism, dependent on indirect interactions among species and non-random spatial organization, in which selection favours restraint in competitive ability. In simple spatial models of a three-species intransitive network, indirect interactions favour slower growth and selection limits the difference in growth rate among species. The mechanism involves a trade-off between selection at the individual level, which selects for increased growth rate, and at the community level, which acts to limit growth rate to less than the maximum possible. If the difference in growth rates among species becomes too large, then the community becomes unstable and collapses to a monoculture of the slowest growing species. The mechanism requires both the intransitive network structure and self-organized spatial structure in the system. Similar behaviours arise in more complex systems of more than three species, and where there are reversals in interaction outcomes between species pairs. The work suggests that spatial self-structuring, indirect interactions and selection acting on community properties can be important in evolution. It provides a partial explanation of the high level of species coexistence and apparent restraint in interspecific interactions evident in some assemblages of sessile marine colonial organisms.

Animals↗

PubMed2002

Sexual selection constrained by life history in a butterfly.

Trait evolution via sexual selection has traditionally been viewed as isolated from life-history constraints. Recent theoretical treatments, however, predict that costly sexually selected characters should be mediated by the same allocational trade-offs that apply to more conventional aspects of reproductive investment. Participation in risky competitive behaviours, for example male-male combat, should therefore increase as the opportunity for future reproduction declines. However, the demonstration of such trade-offs has proven to be elusive due to concomitant age-based variation in the physical determinants of fighting 'ability'. Here, I exploit the unique nature of a butterfly contest system to provide compelling evidence for lifetime partitioning of risky and aggressive sexually selected behaviours. I show that male Hypolimnas bolina become more willing to persist in contests over mating territories, and more generally accepting of injury risks, as they age. Contest persistence in this species is not mediated simply by physical condition, and I experimentally isolate the effect of ageing per se from resource ownership and previous contest experience. These results demonstrate how sexually selected behaviours can be ultimately mediated by a shifting trade-off between contemporary reproductive effort and future opportunities.

Age Factors↗

PubMed2002

Adjustments of ejaculation rates in response to risk of sperm competition in a fish, the bitterling (Rhodeus sericeus).

Game theory models of sperm competition predict that within species, males should increase their sperm expenditure when they have one competitor, but decrease expenditure with increasing numbers of competitors. So far, there have been few tests or support for this prediction. Here, we show that males of a freshwater fish, the European bitterling, Rhodeus sericeus, do indeed adjust their ejaculation rate to the number of male competitors by first increasing and then decreasing their ejaculation rates as the number of competitors increases. However, this occurred only under restricted conditions. Specifically, the prediction was upheld as long as no female had deposited eggs in the live mussels that are used as spawning sites. After one or more females had spawned, males did not decrease their ejaculation rates with the number of competitors, but instead they became more aggressive. This indicates that decreased ejaculation rate and increased aggression are alternative responses to increased risk of sperm competition.

Aggression↗

PubMed2002

Positive fitness consequences of interspecific interaction with a potential competitor.

The coexistence of species sharing mutual resources is usually thought to be limited by negative processes such as interspecific competition. This is because an overlap in resource use leads to negative fitness consequences, and traits favouring avoidance of potential competitors, for example in habitat selection, are therefore selected for. However, species interactions are acknowledged to vary from negative (competition) to mutualism, although empirical evidence for positive interspecific interactions from natural communities of other than plants and sessile animals is scarce. Here, we experimentally examined the habitat selection and its fitness consequences of a migrant bird, the pied flycatcher (Ficedula hypoleuca), in relation to the presence of competitively superior birds, resident titmice (Parus spp.). Experiments were conducted on two spatial scales: landscape and nest-site scale. We demonstrate that pied flycatchers were attracted to and accrued fitness benefits from the presence of titmice. Flycatchers breeding in tight association with titmice initiated breeding earlier, had larger broods and heavier young than solitarily breeding flycatchers. This paradoxical result indicates that species interactions may switch from negative to positive and that the coexistence of species is not always restricted by negative costs caused by other species.

Animals↗

PubMed2002

Procedures for reliable estimation of viral fitness from time-series data.

In order to develop a better understanding of the evolutionary dynamics of HIV drug resistance, it is necessary to quantify accurately the in vivo fitness costs of resistance mutations. However, the reliable estimation of such fitness costs is riddled with both theoretical and experimental difficulties. Experimental fitness assays typically suffer from the shortcoming that they are based on in vitro data. Fitness estimates based on the mathematical analysis of in vivo data, however, are often questionable because the underlying assumptions are not fulfilled. In particular, the assumption that the replication rate of the virus population is constant in time is frequently grossly violated. By extending recent work of Marée and colleagues, we present here a new approach that corrects for time-dependent viral replication in time-series data for growth competition of mutants. This approach allows a reliable estimation of the relative replicative capacity (with confidence intervals) of two competing virus variants growing within the same patient, using longitudinal data for the total plasma virus load, the relative frequency of the two variants and the death rate of infected cells. We assess the accuracy of our method using computer-generated data. An implementation of the developed method is freely accessible on the Web (http://www.eco.ethz.ch/fitness.html).

Anti-HIV Agents↗

PubMed2002

Polarization of competition increases with latitude.

Many organisms overlap in their use of resources in space and time. Where and when resources are restricted, species must compete for them. Living space, often a critical resource controlling food and mate availability, is directly contested by organisms in most habitats. The ensuing animal interactions generally result in a winner gaining space and a loser, which may die. Contact matrices from studies of interference competition in encrusting marine Bryozoa (clonal and colonial animals), spanning at least 60 degrees latitude in both hemispheres, were analysed and subjected to a modern transitivity index. Only data for Bryozoa were used because (i) use of a single taxon with restricted ecology simplifies the scope for types of encounters, (and therefore) interpretation; and (ii) ecological bias is reduced because bryozoans are abundant at all latitudes. The analysis shows that assemblage competition is more hierarchical towards both poles. Thus, poorer competitors fail more frequently in interactions with increasing latitude. The cause of this trend is the simplification of overall outcomes between competitors, such as fewer ties, reversals in outcome or competitive loops (where low-ranking competitors beat those of higher ranking). The implication of such a trend is that the maintenance of biological diversity at high latitudes may principally be by physical rather than biological (competition) processes. Certainly, ocean surface energy increases with latitude through wind and wave action (and ice scour in polar regions).

Animals↗

PubMed2002

Mating system and sex ratios of a pollinating fig wasp with dispersing males.

Recent studies have used sex ratios to quantify the mating systems of organisms, the argument behind it being that more female-biased sex ratios are an indication of higher local mate competition, which goes hand-in-hand with higher levels of inbreeding. Although qualitative tests of the effects of mating systems on sex ratios abound, there is a dearth of studies that quantify both the mating system and the sex ratio. I use a colour dimorphism with a simple Mendelian inheritance to quantify the mating system of an unusual fig-pollinating wasp in which males disperse to obtain matings on non-natal mating patches. In qualitative agreement with initial expectations, the sex ratios of single foundresses are found to be higher than those of regular species. However, by quantifying the mating system, it is shown that the initial expectation is incorrect and this species' sex ratio is a poor predictor of its mating system (it underestimates the frequency of sib-mating). The species has a very high variance in sex ratio suggesting that excess males can simply avoid local mate competition (and hence a lowered fitness to their mother) by dispersing to other patches.

Animals↗

PubMed2002

Competition for male reproductive investment elevates testosterone levels in female dunnocks, Prunella modularis.

In many songbirds, females occasionally sing in contexts of high female-female competition. Testosterone may be involved in the activation of song, because testosterone implants elicit female song in many species with rare female song. A possible mechanism for the hormonal control of female song is provided by the challenge hypothesis, which predicts a rise in testosterone in response to aggressive interactions during socially unstable situations. We tested this by comparing faecal testosterone levels in polygynandrous and monogamous female dunnocks. In groups with two to three females (polygynandry and polygyny) males provide less help at each nest than in groups with a single female (monogamy and polyandry). Polygynandrous and polygynous females are aggressive towards one another and attempt to expel rivals. Polygynandrous females had significantly higher testosterone levels than monogamous females. Competition between females that was induced by removal of males caused testosterone levels to rise. Further, female testosterone levels were correlated with the rate of 'tseep' calls, which are produced during aggressive encounters between females. Finally, polygynandrous and polygynous females sang significantly more than monogamous females. To the best of our knowledge, these results provide the first experimental support for the challenge hypothesis in female birds, and suggest that testosterone can regulate facultative female song in songbirds.

Animals↗

PubMed2002

Interspecific competition during transmission of two sympatric malaria parasite species to the mosquito vector.

The role of species interactions in structuring parasite communities remains controversial. Here, we show that interspecific competition between two avian malaria parasite species, Plasmodium gallinaceum and P. juxtanucleare, occurs as a result of interference during parasite fertilization within the bloodmeal of the mosquito. The significant reduction in the transmission success of P. gallinaceum to mosquitoes, due to the co-infecting P. juxtanucleare, is predicted to have compromised its colonization of regions occupied by P. juxtanucleare and, thus, may have contributed to the restricted global distribution of P. gallinaceum. Such interspecies interactions may occur between human malaria parasites and, thus, impact upon parasite species epidemiology, especially in regions of seasonal transmission.

Aedes↗

PubMed2002

Interference competition and species coexistence.

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

Animals↗