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1.  Host acceptance and sex allocation of Nasonia wasps in response to conspecifics and heterospecifics 
Species recognition is an important aspect of an organism's biology. Here, we consider how parasitoid wasps vary their reproductive decisions when their offspring face intra- and interspecific competition for resources and mates. We use host acceptance and sex ratio behaviour to test whether female Nasonia vitripennis and Nasonia longicornis discriminate between conspecifics and heterospecifics when ovipositing. We tested pairs of conspecific or heterospecific females ovipositing either simultaneously or sequentially on a single host, using strains varying in their recent history of sympatry. Both N. vitripennis and N. longicornis rejected parasitized hosts more often than unparasitized hosts, although females were more likely to superparasitize their own species in the sequential treatment. However, sex ratio behaviour did not vary, suggesting similar responses towards conspecifics and heterospecifics. This contrasts with theory predicting that heterospecifics should not influence sex ratios as their offspring do not influence local mate competition, where conspecifics would. These non-adaptive sex ratios reinforce the lack of adaptive kin discrimination in N. vitripennis and suggest a behavioural constraint. Discrimination between closely related species is therefore context dependent in Nasonia. We suggest that isolating mechanisms associated with the speciation process have influenced behaviour to a greater extent than selection on sex ratios.
doi:10.1098/rspb.2009.0977
PMCID: PMC2817309  PMID: 19640886
species recognition; speciation; adaptation; sex ratios; superparasitism; multiparasitism
2.  High-quality habitat and facilitation ameliorate competitive effects of prior residents on new settlers 
Oecologia  2010;166(1):121-130.
Many species disperse during their lifetime. Two factors that can affect the performance of individuals following dispersal are the presence of conspecifics and intrinsic habitat quality at the settlement site. Detecting the influence of these factors can be difficult for at least two reasons: (1) the outcomes of interactions with conspecifics are often variable including both competition and facilitation, and (2) selection of high quality habitats often leads to positive covariance between habitat quality and density. In this study, I investigate positive and negative effects of resident blue streak cleaner wrasse (Labroides dimidiatus) on the growth and survival of recently settled conspecifics while accounting for habitat quality. Juvenile L. dimidiatus settle near adult conspecifics, but likely have to compete with resident adults for access to food. However, field experiments indicate that settlers have access to more resources at occupied sites, and as a result, grow faster despite evidence for competition with residents. This result is a direct consequence of two factors: (1) resident conspecifics facilitate settlers by attracting client fish, and (2) resident conspecifics are strongly associated with high quality habitat. These results highlight the need to simultaneously consider habitat quality and competitive and facilitative interactions between conspecifics when making inferences about ecological processes from spatial patterns of individual performance.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-010-1826-7) contains supplementary material, which is available to authorized users.
doi:10.1007/s00442-010-1826-7
PMCID: PMC3074066  PMID: 21049298
Density-dependent growth; Habitat selection; Labroides dimidiatus; Coral reef; Fish
3.  Diet and density dependent competition affect larval performance and oviposition site selection in the mosquito species Aedes albopictus (Diptera: Culicidae) 
Parasites & Vectors  2012;5:225.
Background
Oviposition-site choice is an essential component of the life history of all mosquito species. According to the oviposition-preference offspring-performance (P-P) hypothesis, if optimizing offspring performance and fitness ensures high overall reproductive fitness for a given species, the female should accurately assess details of the heterogeneous environment and lay her eggs preferentially in sites with conditions more suitable to offspring.
Methods
We empirically tested the P-P hypothesis using the mosquito species Aedes albopictus by artificially manipulating two habitat conditions: diet (measured as mg of food added to a container) and conspecific density (CD; number of pre-existing larvae of the same species). Immature development (larval mortality, development time to pupation and time to emergence) and fitness (measured as wing length) were monitored from first instar through adult emergence using a factorial experimental design over two ascending gradients of diet (2.0, 3.6, 7.2 and 20 mg food/300 ml water) and CD (0, 20, 40 and 80 larvae/300 ml water). Treatments that exerted the most contrasting values of larval performance were recreated in a second experiment consisting of single-female oviposition site selection assay.
Results
Development time decreased as food concentration increased, except from 7.2 mg to 20.0 mg (Two-Way CR ANOVA Post-Hoc test, P > 0.1). Development time decreased also as conspecific density increased from zero to 80 larvae (Two-Way CR ANOVA Post-Hoc test, P < 0.5). Combined, these results support the role of density-dependent competition for resources as a limiting factor for mosquito larval performance. Oviposition assays indicated that female mosquitoes select for larval habitats with conspecifics and that larval density was more important than diet in driving selection for oviposition sites.
Conclusions
This study supports predictions of the P-P hypothesis and provides a mechanistic understanding of the underlying factors driving mosquito oviposition site selection.
doi:10.1186/1756-3305-5-225
PMCID: PMC3481443  PMID: 23044004
Habitat selection; Oviposition; Diet; Conspecific density; Mosquito
4.  Conspecific and Heterospecific Cues Override Resource Quality to Influence Offspring Production 
PLoS ONE  2013;8(7):e70268.
Animals live in an uncertain world. To reduce uncertainty, animals use cues that can encode diverse information regarding habitat quality, including both non-social and social cues. While it is increasingly appreciated that the sources of potential information are vast, our understanding of how individuals integrate different types of cues to guide decision-making remains limited. We experimentally manipulated both resource quality (presence/absence of cactus fruit) and social cues (conspecific juveniles, heterospecific juveniles, no juveniles) for a cactus-feeding insect, Narniafemorata (Hemiptera: Coreidae), to ask how individuals responded to resource quality in the presence or absence of social cues. Cactus with fruit is a high-quality environment for juvenile development, and indeed we found that females laid 56% more eggs when cactus fruit was present versus when it was absent. However, when conspecific or heterospecific juveniles were present, the effects of resource quality on egg numbers vanished. Overall, N. femorata laid approximately twice as many eggs in the presence of heterospecifics than alone or in the presence of conspecifics. Our results suggest that the presence of both conspecific and heterospecific social cues can disrupt responses of individuals to environmental gradients in resource quality.
doi:10.1371/journal.pone.0070268
PMCID: PMC3704596  PMID: 23861984
5.  Conspecifics as informers and competitors: an experimental study in foraging bumble-bees 
Conspecifics are usually considered competitors negatively affecting food intake rates. However, their presence can also inform about resource quality by providing inadvertent social information. Few studies have investigated whether foragers perceive conspecifics as informers or competitors. Here, we experimentally tested whether variation in the density of demonstrators (‘none’, ‘low’ and ‘high’), whose location indicated flower profitability, affected decision-making of bumble-bees Bombus terrestris. Bumble-bees foraged on either ‘simple’ (two colours) or ‘complex’ (four colours) artificial floral communities. We found that conspecifics at low density may be used as sources of information in first flower choices, whereas they appeared as competitors over the whole foraging sequence. Low conspecific densities improved foragers' first-visit success rate in the simple environment, and decreased time to first landing, especially in the complex environment. High conspecific densities did not affect these behavioural parameters, but reduced flower constancy in both floral communities, which may alter the efficiency of pollinating visits. These results suggest that the balance of the costs and benefits of conspecific presence varies with foraging experience, floral community and density. Spatio-temporal scales could thus be an important determinant of social information use. This behavioural flexibility should allow bumble-bees to better exploit their environment.
doi:10.1098/rspb.2010.2659
PMCID: PMC3145184  PMID: 21288951
Bombus terrestris; decision-making; competition; foraging behaviour; inadvertent social information; plant–pollinator interactions
6.  Hump-Shaped Density-Dependent Regulation of Mosquito Oviposition Site-Selection by Conspecific Immature Stages: Theory, Field Test with Aedes albopictus, and a Meta-Analysis 
PLoS ONE  2014;9(3):e92658.
Oviposition site selection by gravid females is an important determinant of the distribution, abundance, and dynamics of dipteran hematophagous insects. The presence of conspecific immature stages in a potential oviposition site could, on the one hand, indicate the suitability of that site but on the other hand could indicate the potential for intraspecific competition. In this paper, we present a graphic model suggesting that the trade-off between these two opposing forces could result in a hump-shaped density-dependent relationship between oviposition rate and conspecific immature stage density (hereafter, the “Hump-shaped regulation model”) with positive effects of aggregation prevailing at low densities and negative effect of intraspecific competition prevailing at higher densities. We field-tested the predictions of this model at both the egg- and the larval levels with Aedes albopictus and evaluated if and how these relationships are affected by resource enrichment. We found support for the hump-shaped regulation model at both the larval and the egg levels. Using oviposition cups containing varying numbers of conspecific larvae, we showed that the oviposition activity of Ae. albopictus first increases and then decreases with larvae number. Medium enrichment resulted in higher hatching rate, and demonstrated linear relations for the no-enrichment treatment where larvae density range was low and hump-shaped relationship for the enriched medium that had a wider larvae density range. Using pairs of oviposition cups, we showed that at low egg densities mosquitoes laid more eggs on substrates containing pre-existing eggs. However, at higher egg densities, mosquitoes laid more eggs on a virgin substrate. Based on our results and on a meta-analysis, we suggest that due to study design or methodological shortcomings the hump-shaped regulation model is often left undetected and that it is likely to be more common than currently thought.
doi:10.1371/journal.pone.0092658
PMCID: PMC3969363  PMID: 24681526
7.  Adaptive significance of permanent female mimicry in a bird of prey 
Biology Letters  2011;8(2):167-170.
Permanent female mimicry, in which adult males express a female phenotype, is known only from two bird species. A likely benefit of female mimicry is reduced intrasexual competition, allowing female-like males to access breeding resources while avoiding costly fights with typical territorial males. We tested this hypothesis in a population of marsh harriers Circus aeruginosus in which approximately 40 per cent of sexually mature males exhibit a permanent, i.e. lifelong, female plumage phenotype. Using simulated territorial intrusions, we measured aggressive responses of breeding males towards conspecific decoys of females, female-like males and typical males. We show that aggressive responses varied with both the type of decoys and the type of defending male. Typical males were aggressive towards typical male decoys more than they were towards female-like male decoys; female-like male decoys were attacked at a rate similar to that of female decoys. By contrast, female-like males tolerated male decoys (both typical and female-like) and directed their aggression towards female decoys. Thus, agonistic responses were intrasexual in typical males but intersexual in female-like males, indicating that the latter not only look like females but also behave like them when defending breeding resources. When intrasexual aggression is high, permanent female mimicry is arguably adaptive and could be seen as a permanent ‘non-aggression pact’ with other males.
doi:10.1098/rsbl.2011.0914
PMCID: PMC3297406  PMID: 22072281
colour morph; alternative behavioural strategies; paternity assurance; marsh harrier; sexual dichromatism
8.  Breeding state and season affect interspecific interaction types: indirect resource competition and direct interference 
Oecologia  2011;167(3):623-633.
Indirect resource competition and interference are widely occurring mechanisms of interspecific interactions. We have studied the seasonal expression of these two interaction types within a two-species, boreal small mammal system. Seasons differ by resource availability, individual breeding state and intraspecific social system. Live-trapping methods were used to monitor space use and reproduction in 14 experimental populations of bank voles Myodes glareolus in large outdoor enclosures with and without a dominant competitor, the field vole Microtus agrestis. We further compared vole behaviour using staged dyadic encounters in neutral arenas in both seasons. Survival of the non-breeding overwintering bank voles was not affected by competition. In the spring, the numbers of male bank voles, but not of females, were reduced significantly in the competition populations. Bank vole home ranges expanded with vole density in the presence of competitors, indicating food limitation. A comparison of behaviour between seasons based on an analysis of similarity revealed an avoidance of costly aggression against opponents, independent of species. Interactions were more aggressive during the summer than during the winter, and heterospecific encounters were more aggressive than conspecific encounters. Based on these results, we suggest that interaction types and their respective mechanisms are not either–or categories and may change over the seasons. During the winter, energy constraints and thermoregulatory needs decrease direct aggression, but food constraints increase indirect resource competition. Direct interference appears in the summer, probably triggered by each individual’s reproductive and hormonal state and the defence of offspring against conspecific and heterospecific intruders. Both interaction forms overlap in the spring, possibly contributing to spring declines in the numbers of subordinate species.
doi:10.1007/s00442-011-2008-y
PMCID: PMC3193992  PMID: 21597944
Rodents; Aggression; Seasonality; Space use; Winter biology
9.  Differential behavioural and endocrine responses of common voles (Microtus arvalis) to nest predators and resource competitors 
BMC Ecology  2013;13:33.
Background
Adaptive behavioural strategies promoting co-occurrence of competing species are known to result from a sympatric evolutionary past. Strategies should be different for indirect resource competition (exploitation, e.g., foraging and avoidance behaviour) than for direct interspecific interference (e.g., aggression, vigilance, and nest guarding). We studied the effects of resource competition and nest predation in sympatric small mammal species using semi-fossorial voles and shrews, which prey on vole offspring during their sensitive nestling phase. Experiments were conducted in caged outdoor enclosures. Focus common vole mothers (Microtus arvalis) were either caged with a greater white-toothed shrew (Crocidura russula) as a potential nest predator, with an herbivorous field vole (Microtus agrestis) as a heterospecific resource competitor, or with a conspecific resource competitor.
Results
We studied behavioural adaptations of vole mothers during pregnancy, parturition, and early lactation, specifically modifications of the burrow architecture and activity at burrow entrances. Further, we measured pre- and postpartum faecal corticosterone metabolites (FCMs) of mothers to test for elevated stress hormone levels. Only in the presence of the nest predator were prepartum FCMs elevated, but we found no loss of vole nestlings and no differences in nestling body weight in the presence of the nest predator or the heterospecific resource competitor. Although the presence of both the shrew and the field vole induced prepartum modifications to the burrow architecture, only nest predators caused an increase in vigilance time at burrow entrances during the sensitive nestling phase.
Conclusion
Voles displayed an adequate behavioural response for both resource competitors and nest predators. They modified burrow architecture to improve nest guarding and increased their vigilance at burrow entrances to enhance offspring survival chances. Our study revealed differential behavioural adaptations to resource competitors and nest predators.
doi:10.1186/1472-6785-13-33
PMCID: PMC3847765  PMID: 24010574
Behavioural adaptations; Small mammals; Interspecific interactions; Nest predation; Stress response; Faecal corticosterone metabolites; Burrow system; Shrews; Voles
10.  Oviposition Site Selection by the Dengue Vector Aedes aegypti and Its Implications for Dengue Control 
Background
Because no dengue vaccine or antiviral therapy is commercially available, controlling the primary mosquito vector, Aedes aegypti, is currently the only means to prevent dengue outbreaks. Traditional models of Ae. aegypti assume that population dynamics are regulated by density-dependent larval competition for food and little affected by oviposition behavior. Due to direct impacts on offspring survival and development, however, mosquito choice in oviposition site can have important consequences for population regulation that should be taken into account when designing vector control programs.
Methodology/Principal Findings
We examined oviposition patterns by Ae. aegypti among 591 naturally occurring containers and a set of experimental containers in Iquitos, Peru. Using larval starvation bioassays as an indirect measure of container food content, we assessed whether females select containers with the most food for their offspring. Our data indicate that choice of egg-laying site is influenced by conspecific larvae and pupae, container fill method, container size, lid, and sun exposure. Although larval food positively influenced oviposition, our results did not support the hypothesis that females act primarily to maximize food for larvae. Females were most strongly attracted to sites containing immature conspecifics, even when potential competitors for their progeny were present in abundance.
Conclusion/Significance
Due to strong conspecific attraction, egg-laying behavior may contribute more to regulating Ae. aegypti populations than previously thought. If highly infested containers are targeted for removal or larvicide application, females that would have preferentially oviposited in those sites may instead distribute their eggs among other suitable, previously unoccupied containers. Strategies that kill mosquitoes late in their development (i.e., insect growth regulators that kill pupae rather than larvae) will enhance vector control by creating “egg sinks,” treated sites that exploit conspecific attraction of ovipositing females, but reduce emergence of adult mosquitoes via density-dependent larval competition and late acting insecticide.
Author Summary
Controlling the mosquito Aedes aegypti is of public health importance because, at present, it is the only means to stop dengue virus transmission. Implementing successful mosquito control programs requires understanding what factors regulate population abundance, as well as anticipating how mosquitoes may adapt to control measures. In some species of mosquitoes, females choose egg-laying sites to improve the survival and growth of their offspring, a behavior that ultimately influences population distribution and abundance. In the current study, we tested whether Ae. aegypti actively choose the containers in which they lay their eggs and determined what cues are most relevant to that process. We also explored whether females select containers that provide the most food for their larval progeny. Surprisingly, egg-laying females were most attracted to sites containing other immature Ae. aegypti, rather than to sites containing the most food. We propose that this behavior may contribute to density-dependent competition for food among larvae and play a larger role than previously thought in regulating Ae. aegypti populations. We recommend that accounting for, and even taking advantage of, this natural behavior will lead to more effective strategies for dengue prevention.
doi:10.1371/journal.pntd.0001015
PMCID: PMC3075222  PMID: 21532736
11.  Experimental evidence for asymmetric mate preference and aggression: behavioral interactions in a woodrat (Neotoma) hybrid zone 
Background
Female mate preferences may be under strong selection in zones of contact between closely related species because of greater variation in available mates and the potential costs of hybridization. We studied female mate preferences experimentally in a zone of secondary contact between Desert and Bryant’s Woodrat (Neotoma lepida and N. bryanti) in the southern foothills of the Sierra Nevada of California. We tested female preference for conspecific versus heterospecific males in paired choice trials in which females could interact freely with males, but males could not interact directly with each other. We compared preferences of females from both allopatric and sympatric sites.
Results
We did not find evidence of the process of reinforcement as assortative preferences were not stronger in sympatry than in allopatry. Mate preferences, however, were asymmetric, with N. lepida females mating preferentially with conspecifics and N. bryanti females showing no preference by species. Sympatric females were less likely to mate than allopatric females, due in part to an increase in aggressive interactions. However, even in the absence of aggression, courtship led to mating less often in sympatric females, suggesting they were choosier or had lower sexual motivation than allopatric females.
Conclusions
Patterns of mate choice in this woodrat system appear to be strongly impacted by body size and aggressive behavior. In particular, females of the smaller-bodied species rarely interact with the relatively large heterospecific males. In contrast females of the larger-bodied species accept the relatively small heterospecific males. For sympatric animals, rates of aggression were markedly higher than for allopatric animals and reduced affiliative and reproductive behavior in our trials. Sympatric animals are larger and more aggressive, traits that are likely under strong ecological selection across the sharp resource gradient that characterizes the contact zone. However, our results suggest that these traits that are likely favored in competitive interactions between the species also impact reproductive interactions. Combined with our previous findings of post-zygotic isolation in this system, this study suggests that multiple isolating mechanisms contribute to the rate of genetic exchange between these species when they come into contact, and that these mechanisms are the result of selection on traits that are important in a range of ecological and reproductive interactions.
doi:10.1186/1471-2148-13-220
PMCID: PMC3853547  PMID: 24093823
Hybridization; Reproductive isolation; Reproductive character displacement; Reinforcement
12.  Female Sexual Arousal in Amphibians 
Hormones and behavior  2010;59(5):630-636.
Rather than being a static, species specific trait, reproductive behavior in female amphibians is variable within an individual during the breeding season when females are capable of reproductive activity. Changes in receptivity coincide with changes in circulating estrogen. Estrogen is highest at the point when females are ready to choose a male and lay eggs. At this time female receptivity (her probability of responding to a male vocal signal) is highest and her selectivity among conspecific calls (measured by her probability of responding to a degraded or otherwise usually unattractive male signal) is lowest. These changes occur even though females retain the ability to discriminate different acoustic characteristics of various conspecific calls. After releasing her eggs, female amphibians quickly become less receptive and more choosy in terms of their responses to male sexual advertisement signals. Male vocal signals stimulate both behavior and estrogen changes in amphibian females making mating more probable. The changes in female reproductive behavior are the same as those generally accepted as indicative of a change in female sexual arousal leading to copulation. They are situationally triggered, gated by interactions with males, and decline with the consummation of sexual reproduction with a chosen male. The changes can be triggered by either internal physiological state or by the presence of stimuli presented by males, and the same stimuli change both behavior and physiological (endocrine) state in such a way as to make acceptance of a male more likely. Thus amphibian females demonstrate many of the same general characteristics of changing female sexual state that in mammals indicate sexual arousal.
doi:10.1016/j.yhbeh.2010.08.015
PMCID: PMC3008597  PMID: 20816968
amphibians; reproduction; female; communication; estrogen
13.  The biology and behavior of the longhorned beetle, Dectes texanus on sunflower and soybean 
The biology and behavior of the longhorned beetle Dectes texanus LeConte (Coleoptera: Cerambycidae) was studied on two host plants that suffer economic losses from this pest; sunflower, Helianthus annuus, and soybean, Glycines max. Reciprocal crosses of D. texanus collected from the two plants all produced viable progeny, indicating that conspecific insects attack both crops. Pupae from soybean stalks weighed about 40% less than those from sunflower, and adults fed on soybean lived a mean of 23 days, compared to a mean of 53 days (males) and 76 days (females) for those fed sunflower. A female's larval host plant had no effect on her tendency to ovipuncture plants of either type in a greenhouse trial. A field-tested population collected exclusively from sunflower contained three types of females in similar proportions: those that laid eggs only on sunflower, those that laid only on soybean, and those that laid equally on both host plants. Females in field trials fed more on the plant they had fed on in the laboratory, but soybean-fed females fed more on soybean than did sunflower-fed females. Females fed soybean also made more ovipunctures on soybean plants in field trials than sunflower-fed females, but their responses to sunflower plants were similar. Females displayed higher total ovipositional activity when they encountered sunflower first in the field, and lower total activity when they encountered soybean first. Feeding scores were significantly correlated with ovipunctures and eggs on both plant types. We conclude that sunflower is the preferred host plant, although females will accept soybean when it is the only available food. The results suggest that D. texanus is still in the initial stages of a host range expansion with female host selection behavior demonstrating both genetic influences and phenotypic flexibility. Sunflower represents a nutritionally superior, ancestral host plant and relatively high fitness costs are still associated with utilization of the novel host plant, soybean, costs that may be offset by benefits such as reduced intraspecific competition. These potential benefits and their consequent implications for D. texanus host range evolution are hypothesized and discussed.
PMCID: PMC1615232  PMID: 17119607
evolution; fitness; Glycines max; Helianthus annuus; host range; longevity; oviposition; polyphagy; reproduction
14.  Costs of Defense and a Test of the Carbon-Nutrient Balance and Growth-Differentiation Balance Hypotheses for Two Co-Occurring Classes of Plant Defense 
PLoS ONE  2012;7(10):e47554.
One of the goals of chemical ecology is to assess costs of plant defenses. Intraspecific trade-offs between growth and defense are traditionally viewed in the context of the carbon-nutrient balance hypothesis (CNBH) and the growth-differentiation balance hypothesis (GDBH). Broadly, these hypotheses suggest that growth is limited by deficiencies in carbon or nitrogen while rates of photosynthesis remain unchanged, and the subsequent reduced growth results in the more abundant resource being invested in increased defense (mass-balance based allocation). The GDBH further predicts trade-offs in growth and defense should only be observed when resources are abundant. Most support for these hypotheses comes from work with phenolics. We examined trade-offs related to production of two classes of defenses, saponins (triterpenoids) and flavans (phenolics), in Pentaclethra macroloba (Fabaceae), an abundant tree in Costa Rican wet forests. We quantified physiological costs of plant defenses by measuring photosynthetic parameters (which are often assumed to be stable) in addition to biomass. Pentaclethra macroloba were grown in full sunlight or shade under three levels of nitrogen alone or with conspecific neighbors that could potentially alter nutrient availability via competition or facilitation. Biomass and photosynthesis were not affected by nitrogen or competition for seedlings in full sunlight, but they responded positively to nitrogen in shade-grown plants. The trade-off predicted by the GDBH between growth and metabolite production was only present between flavans and biomass in sun-grown plants (abundant resource conditions). Support was also only partial for the CNBH as flavans declined with nitrogen but saponins increased. This suggests saponin production should be considered in terms of detailed biosynthetic pathway models while phenolic production fits mass-balance based allocation models (such as the CNBH). Contrary to expectations based on the two defense hypotheses, trade-offs were found between defenses and photosynthesis, indicating that studies of plant defenses should include direct measures of physiological responses.
doi:10.1371/journal.pone.0047554
PMCID: PMC3480385  PMID: 23115654
15.  When Violence Pays: A Cost-Benefit Analysis of Aggressive Behavior in Animals and Humans 
An optimization analysis of human behavior from a comparative perspective can improve our understanding of the adaptiveness of human nature. Intra-specific competition for resources provides the main selective pressure for the evolution of violent aggression toward conspecifics, and variation in the fitness benefits and costs of aggression can account for inter-specific and inter-individual differences in aggressiveness. When aggression reflects competition for resources, its benefits vary in relation to the characteristics of the resources (their intrinsic value, abundance, spatial distribution, and controllability) while its costs vary in relation to the characteristics of organisms and how they fight (which, in turn, affects the extent to which aggression entails risk of physical injury or death, energetic depletion, exposure to predation, psychological and physiological stress, or damage to social relationships). Humans are a highly aggressive species in comparison to other animals, probably as a result of an unusually high benefit-to-cost ratio for intra-specific aggression. This conclusion is supported by frequent and widespread occurrence of male-male coalitionary killing and by male-female sexual coercion. Sex differences in violent aggression in humans and other species probably evolved by sexual selection and reflect different optimal competitive strategies for males and females.
PMCID: PMC3859192  PMID: 23864299
aggression; evolution; optimization; competition; violence
16.  Scramble in behaviour and ecology. 
Nicholson's distinction between 'scramble' and 'contest' modes of competition has received widespread attention in ecology and in behaviour, though the emphasis has been different between the two disciplines. In ecology the focus has been on the effects on population; in behavioural ecology the focus has been on the consequences at the individual level. This paper reviews and develops a theory of scramble competition at the individual level, deriving a general evolutionarily stable strategy (ESS) for individual scramble expenditure in a patchy habitat in which individuals compete in local groups for available resources, and examines two population consequences. The critical parameter determining the relationship between individual scramble expenditure and the number of competitors in a patch is the expected resource per capita. If resource input, R, to a patch is constant and independent of the number of competitors, n, then as the number of competitors increases, the per-capita resources declines as R/n, and the ESS scramble level declines (in proportion to (n-1)/n2). However, if the resource input to a patch is positively related to the number of competitors in the patch, scramble expenditure may increase with the number of competitors. In the case where the per-capita resource input stays constant (i.e. R(n) = Rn), the scramble level increases with competitor number (in proportion to (n-1) /n). There are plausible ecological reasons why either of these extreme limits may be approached in nature, making it important to ascertain the relationship between R and n before predicting individual scramble expenditure. For example, resource input may be constant when groups of competitors are constrained to remain together in given patches, and constant per-capita resources may be approached when ideal-free foraging rules apply. However, in the latter case, scramble expenditure must be accounted for in determining the ideal-free distribution. An analysis shows that this leads to 'undermatching', i.e. the ratio of numbers of competitors for good/bad patches becomes progressively less than the ratio of input rates for good/bad patches as the difference between the good and bad patches increases. A second population consequence of the scramble ESS relates to the fact that scrambles may dramatically affect fitness. The per-capita gain in energy can be reduced by a factor of up to 1/n as a result of scramble expenditure, potentially reducing realized population size to as little as the square root of the maximum potential carrying capacity, though reasons are given why such large reductions are unlikely.
PMCID: PMC1692901  PMID: 11127910
17.  Unexpected male choosiness for mates in a spider 
Sexual selection theory traditionally considers choosiness for mates to be negatively related to intra-sexual competition. Males were classically considered to be the competing, but not the choosy, sex. However, evidence of male choosiness is now accumulating. Male choosiness is expected to increase with an individual's competitive ability, and to decrease as intra-sexual competition increases. However, such predictions have never been tested in field conditions. Here, we explore male mate choice in a spider by studying size-assortative pairing in two natural sites that strongly differ in the level of male–male competition. Unexpectedly, our results demonstrate that mate choice shifts from opportunism to high selectivity as competition between males increases. Males experiencing weak competition did not exhibit size-related mating preferences. By contrast, when competition was intense we found strong size-assortative pairing due to male choice: while larger, more competitive males preferentially paired with larger, more fecund females, smaller males chose smaller females. Thus, we show that mating preferences of males vary with their competitive ability. The distinct preferences exhibited by males of different sizes seem to be an adaptive response to the lower reproductive opportunities arising from increased competition between males.
doi:10.1098/rspb.2007.1278
PMCID: PMC2562406  PMID: 17956845
sexual selection; male mate choice; intra-sexual competition; assortative mating; spider
18.  Body condition but not dietary restriction prolongs lifespan in a semelparous capital breeder 
Biology Letters  2009;5(5):636-638.
Effects of diet on longevity are complex because acquired resources are shared among growth, reproduction and somatic maintenance. We simplify these axes by examining how dietary restriction and competitive contexts affect longevity using semelparous males of the Australian redback spider (Latrodectus hasselti). Plastic development of L. hasselti males results in trade-offs of body condition against faster development if females are present, facilitating scramble competition. In the absence of females, males develop slowly as high body condition adults, and are better equipped for mate searching. Here we focus on effects of diet and competitive context on body condition and longevity. Although male survival depended on body condition and exercise, contrary to studies in a wide range of taxa, dietary restriction did not increase longevity. However, there was an interactive effect of diet and competitive context on lifespan, because high-diet males reared in the absence of females lived longer than males reared in the presence of females. Thus males near females pay a survival cost of developing rapidly. This shows that life-history trade-offs affected by competitive context can impose longevity costs independent of the direct energy expenditure of searching, courtship, competition or reproduction.
doi:10.1098/rsbl.2009.0335
PMCID: PMC2781961  PMID: 19515652
dietary restriction; life-history trade-offs; longevity; developmental plasticity; Latrodectus hasselti
19.  Competition for food in a solitarily foraging folivorous primate (Lepilemur leucopus)? 
American Journal of Primatology  2014;76(9):842-854.
Group-living folivorous primates can experience competition for food, and feeding competition has also been documented for solitarily foraging gummivorous and omnivorous primates. However, little is known about the types and consequences of feeding competition in solitary folivorous foragers. We conducted this study in the spiny forest of Berenty Reserve, southern Madagascar, to characterize the competitive regime of the nocturnal solitarily foraging white-footed sportive lemur (Lepilemur leucopus), a species that lives in dispersed pairs. We analyzed 1,213 hr of behavioral observations recorded simultaneously for the male and female of each of seven social units and recorded seasonal changes in food availability over a complete annual cycle. Lepilemur leucopus exhibited low selectivity in its dietary choice and mainly included the most abundant plant species in its diet. Contrary to our predictions, we did not find evidence for increased rates of contest (i.e., displacement from food trees) or scramble competition (i.e., shared use of food patches) during the lean season, neither within nor between social units. Instead, conflict rates were low throughout the year, and, during these observations, any feeding stress may have been more related to food quality than quantity. The resource defense hypotheses may not explain pair-living in this species as there was no indication that males defend food resources for their female pair-partners. The observed lack of feeding competition may indicate that a cryptic anti-predator strategy is a better predictor of spatial avoidance of pair-partners than conflict over food. While anti-predator benefits of crypsis may explain, at least partly, female-female avoidance, studies on the relationship between territory size/quality and reproductive success are required to understand whether feeding competition reduces the potential for female association in L. leucopus.
doi:10.1002/ajp.22272
PMCID: PMC4229058  PMID: 24677259
feeding competition; resource defense; seasonality; spatial avoidance; predation; crypsis
20.  Proximate Factors Underpinning Receiver Responses to Deceptive False Alarm Calls in Wild Tufted Capuchin Monkeys: Is It Counterdeception? 
American Journal of Primatology  2012;75(7):715-725.
Previous research demonstrates that tufted capuchin monkeys use terrestrial predator alarm calls in a functionally deceptive manner to distract conspecifics when feeding on contestable resources, although the success of this tactic is limited because listeners frequently ignore these calls when given in such situations. While this decreased response rate is suggestive of a counterstrategy to deception by receivers, the proximate factors underpinning the behavior are unclear. The current study aims to test if the decreased response rate to alarm calls in competitive contexts is better explained by the perception of subtle acoustic differences between predator-elicited and deceptive false alarms, or by receivers varying their responses based on the context in which the signal is received. This was tested by first examining the acoustic structure of predator-elicited and deceptive false alarms for any potentially perceptible acoustic differences, and second by comparing the responses of capuchins to playbacks of each of predator-elicited and false alarms, played back in noncompetitive contexts. The results indicate that deceptive false alarms and predator-elicited alarms show, at best, minimal acoustic differences based on the structural features measured. Likewise, playbacks of deceptive false alarms elicited antipredator reactions at the same rate as did predator-elicited alarms, although there was a nonsignificant tendency for false alarms to be more likely to elicit escape reactions. The lack of robust acoustic differences together with the high response rate to false alarms in noncompetitive contexts suggests that the context in which the signal is received best explains receiver responses. It remains unclear, however, if listeners ascribe different meanings to the calls based on context, or if they generally ignore all signals in competitive contexts. Whether or not the decreased response rate of receivers directly stems from the deceptive use of the calls cannot be determined until these latter possibilities are rigorously tested. Am. J. Primatol. 75:715-725, 2013. © 2012 Wiley Periodicals, Inc.
doi:10.1002/ajp.22097
PMCID: PMC3746121  PMID: 23184736
vocal communication; deception; skeptical responding; signal reliability; pragmatics; playback experiments
21.  Pheromone Diversification and Age-Dependent Behavioural Plasticity Decrease Interspecific Mating Costs in Nasonia 
PLoS ONE  2014;9(2):e89214.
Interspecific mating can cause severe fitness costs due to the fact that hybrids are often non-viable or less fit. Thus, theory predicts the selection of traits that lessen reproductive interactions between closely related sympatric species. Males of the parasitic wasp Nasonia vitripennis differ from all other Nasonia species by an additional sex pheromone component, but the ecological selective forces underlying this pheromone diversification are unknown. Here we present data from lab experiments suggesting that costly interspecific sexual interactions with the sympatric species N. giraulti might have been responsible for the pheromone evolution and some courtship-related behavioural adaptations in N. vitripennis. Most N. giraulti females are inseminated already within the host, but N. giraulti males still invest in costly sex pheromones after emergence. Furthermore, they do not discriminate between N. vitripennis females and conspecifics during courtship. Therefore, N. vitripennis females, most of which emerge as virgins, face the risk of mating with N. giraulti resulting in costly all-male broods due to Wolbachia-induced cytoplasmic incompatibility. As a counter adaptation, young N. vitripennis females discriminate against N. giraulti males using the more complex conspecific sex pheromone and reject most of them during courtship. With increasing age, however, N. vitripennis females become less choosy, but often compensate mating errors by re-mating with a conspecific. By doing so, they can principally avoid suboptimal offspring sex ratios, but a microcosm experiment suggests that under more natural conditions N. vitripennis females cannot completely avoid fitness costs due to heterospecific mating. Our study provides support for the hypothesis that communication interference of closely related sympatric species using similar sexual signals can generate selective pressures that lead to their divergence.
doi:10.1371/journal.pone.0089214
PMCID: PMC3925242  PMID: 24551238
22.  The early social environment affects social competence in a cooperative breeder 
Animal Behaviour  2012;83(4):1067-1074.
Social competence is defined as the ability of an animal to optimize the expression of social behaviour as a function of the available social information. The social environment encountered early in life can affect the expression of various social behaviours later in life. We investigated whether early social experience can affect social competence. In the cooperatively breeding cichlid Neolamprologus pulcher, we tested whether individuals reared with older brood-caring conspecifics persistently perform better in a series of tasks (1) simulating different social contexts, (2) assigning individuals different social roles and (3) exposing them to an unknown social situation. Fish that had been reared together with older conspecifics showed more appropriate behaviours both as winners (more aggressive displays) and as losers (more submissive displays) when aggressively competing with peers over a resource, and when trying to be accepted as subordinate group member and prospective brood care helper by an unfamiliar dominant pair (more submissive displays near shelters), a situation they had never encountered before. In both tasks fish that had grown up with older fish were tolerated better by conspecifics than fish reared with same-age siblings only. We detected effects of the early environment on social behaviour in the juvenile and adult stages of the test fish. Our results suggest that growing up in more complex social groups fosters a general social ability (i.e. social competence) in N. pulcher that improves their performance across different social roles and contexts, and which may provide fitness benefits.
Highlights
► We tested the effects of the early social environment on social competence in fish. ► A complex early social environment induced more appropriate social behaviour. ► Social competence differed across social roles and social contexts. ► The early-environment effects persisted until the adult period. ► Our results suggest that a better social competence yields fitness benefits.
doi:10.1016/j.anbehav.2012.01.037
PMCID: PMC3328768  PMID: 22536004
aggressive behaviour; cichlid; cooperative breeder; development; early environment; Neolamprologus pulcher; social competence; social experience; social skill; submissive behaviour
23.  How Food Controls Aggression in Drosophila 
PLoS ONE  2014;9(8):e105626.
How animals use sensory information to weigh the risks vs. benefits of behavioral decisions remains poorly understood. Inter-male aggression is triggered when animals perceive both the presence of an appetitive resource, such as food or females, and of competing conspecific males. How such signals are detected and integrated to control the decision to fight is not clear. For instance, it is unclear whether food increases aggression directly, or as a secondary consequence of increased social interactions caused by attraction to food. Here we use the vinegar fly, Drosophila melanogaster, to investigate the manner by which food influences aggression. We show that food promotes aggression in flies, and that it does so independently of any effect on frequency of contact between males, increase in locomotor activity or general enhancement of social interactions. Importantly, the level of aggression depends on the absolute amount of food, rather than on its surface area or concentration. When food resources exceed a certain level, aggression is diminished, suggestive of reduced competition. Finally, we show that detection of sugar via Gr5a+ gustatory receptor neurons (GRNs) is necessary for food-promoted aggression. These data demonstrate that food exerts a specific effect to promote aggression in male flies, and that this effect is mediated, at least in part, by sweet-sensing GRNs.
doi:10.1371/journal.pone.0105626
PMCID: PMC4146546  PMID: 25162609
24.  Feeding-induced rearrangement of green leaf volatiles reduces moth oviposition 
eLife  2013;2:e00421.
The ability to decrypt volatile plant signals is essential if herbivorous insects are to optimize their choice of host plants for their offspring. Green leaf volatiles (GLVs) constitute a widespread group of defensive plant volatiles that convey a herbivory-specific message via their isomeric composition: feeding of the tobacco hornworm Manduca sexta converts (Z)-3- to (E)-2-GLVs thereby attracting predatory insects. Here we show that this isomer-coded message is monitored by ovipositing M. sexta females. We detected the isomeric shift in the host plant Datura wrightii and performed functional imaging in the primary olfactory center of M. sexta females with GLV structural isomers. We identified two isomer-specific regions responding to either (Z)-3- or (E)-2-hexenyl acetate. Field experiments demonstrated that ovipositing Manduca moths preferred (Z)-3-perfumed D. wrightii over (E)-2-perfumed plants. These results show that (E)-2-GLVs and/or specific (Z)-3/(E)-2-ratios provide information regarding host plant attack by conspecifics that ovipositing hawkmoths use for host plant selection.
DOI: http://dx.doi.org/10.7554/eLife.00421.001
eLife digest
Plants have developed a variety of strategies to defend themselves against herbivorous animals, particularly insects. In addition to mechanical defences such as thorns and spines, plants also produce compounds known as secondary metabolites that keep insects and other herbivores at bay by acting as repellents or toxins. Some of these metabolites are produced on a continuous basis by plants, whereas others—notably compounds called green-leaf volatiles—are only produced once the plant has been attacked. Green-leaf volatiles—which are also responsible for the smell of freshly cut grass—have been observed to provide plants with both direct protection, by inhibiting or repelling herbivores, and indirect protection, by attracting predators of the herbivores themselves.
The hawkmoth Manduca sexta lays its eggs on various plants, including tobacco plants and sacred Datura plants. Once the eggs have hatched into caterpillars, they start eating the leaves of their host plant, and if present in large numbers, these caterpillars can quickly defoliate and destroy it. In an effort to defend itself, the host plant releases green-leaf volatiles to attract various species of Geocoris, and these bugs eat the eggs.
One of the green-leaf volatiles released by tobacco plants is known as (Z)-3-hexenal, but enzymes released by M. sexta caterpillars change some of these molecules into (E)-2-hexenal, which has the same chemical formula but a different structure. The resulting changes in the ‘volatile profile’ alerts Geocoris bugs to the presence of M. sexta eggs and caterpillars on the plant.
Now Allmann et al. show that adult female M. sexta moths can also detect similar changes in the volatile profile emitted by sacred Datura plants that have been damaged by M. sexta caterpillars. This alerts the moths to the fact that Geocoris bugs are likely to be attacking eggs and caterpillars on the plant, or on their way to the plant, so they lay their eggs on other plants. This reduces competition for resources and also reduces the risk of newly laid eggs being eaten by predators. Allmann et al. also identified the neural mechanism that allows moths to detect changes in the volatile profile of plants—the E- and Z- odours lead to different activation patterns in the moth brain.
DOI: http://dx.doi.org/10.7554/eLife.00421.002
doi:10.7554/eLife.00421
PMCID: PMC3654435  PMID: 23682312
Manduca sexta; plant volatiles; oviposition; Ca imaging; Datura wrightii; Other
25.  Sperm investment in male meadow voles is affected by the condition of the nearby male conspecifics 
Behavioral Ecology  2008;19(6):1159-1164.
Sperm competition occurs when 2 or more males copulate with a particular female during the same reproductive cycle, and their sperm compete to fertilize the female's available eggs. One strategy that male voles use to assess the risk and intensity of sperm competition involves responding to the presence of scent marks of conspecific males found near a sexually receptive female. Previously, we have shown that if a male vole copulated with a female while he was in the presence of the odors of another male he increased his sperm investment relative to his investment if another male's odors were not present. The aim of the present study was to test the hypothesis that males assess differences in the relative quality of competing males and adjust their sperm investment accordingly. We did so by allowing males to copulate when they were exposed to the scent mark of a 24-h food-deprived male (low-quality male) or the scent mark of a male that was not food deprived (high-quality male). The data indicate that male meadow voles did not increase their sperm investment during copulation when exposed to the scent mark of a food-deprived male but did so when they were exposed to the scent mark of a male that was not food deprived. The results support the hypothesis that male voles are able to adjust sperm investment when they encounter the scent marks of males that differ in quality.
doi:10.1093/beheco/arn092
PMCID: PMC2583107  PMID: 19529815
chemical signals; copulatory behavior; food deprivation; scent marking; sperm competition; voles

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