Substrate temperature constrains recruitment and trail following behavior in ants.

In many ant species, foragers use pheromones to communicate the location of resources to nestmates. Mass-recruiting species deposit long-lasting anonymous chemical trails, while group-recruiting species use temporary chemical trails. We studied how high temperature influenced the foraging behavior of a mass-recruiting species (Tapinoma nigerrimum) and a group-recruiting species (Aphaenogaster senilis) through pheromone decay. First, under controlled laboratory conditions, we examined the effect of temperature on the trail pheromone of both species. A substrate, simulating soil, marked with gaster extract was heated for 10 min. at 25°, 35°, 45°, or 55 °C and offered to workers in a choice test. Heating gaster extract reduced the trail following behavior of the mass-recruiters significantly more than that of the group-recruiters. Second, analyses of the chemicals present on the substrate indicated that most T. nigerrimum gaster secretions vanished at 25 °C, and only iridodials persisted up to 55 °C. By contrast, A. senilis secretions were less volatile and resisted better to elevated temperatures to some extent. However, at 55 °C, the only chemicals that persisted were nonadecene and nonadecane. Overall, our results suggest that the foraging behavior of the group-recruiting species A. senilis is less affected by pheromone evaporation than that of the mass-recruiting species T. nigerrimum. This group-recruiting species might, thus, be particularly adapted to environments with fluctuating temperatures.

Temperature limits trail following behaviour through pheromone decay in ants.

In Mediterranean habitats, temperature affects both ant foraging behaviour and community structure. Many studies have shown that dominant species often forage at lower temperature than subordinates. Yet, the factors that constrain dominant species foraging activity in hot environments are still elusive. We used the dominant ant Tapinoma nigerrimum as a model species to test the hypothesis that high temperatures hinder trail following behaviour by accelerating pheromone degradation. First, field observations showed that high temperatures (> 30°C) reduce the foraging activity of T. nigerrimum independently of the daily and seasonal rhythms of this species. Second, we isolated the effect of high temperatures on pheromone trail efficacy from its effect on worker physiology. A marked substrate was heated during 10 min (five temperature treatments from 25°C to 60°C), cooled down to 25°C, and offered in a test choice to workers. At hot temperature treatments (>40°C), workers did not discriminate the previously marked substrate. High temperatures appeared therefore to accelerate pheromone degradation. Third, we assessed the pheromone decay dynamics by a mechanistic model fitted with Bayesian inference. The model predicted ant choice through the evolution of pheromone concentration on trails as a function of both temperature and time since pheromone deposition. Overall, our results highlighted that the effect of high temperatures on recruitment intensity was partly due to pheromone evaporation. In the Mediterranean ant communities, this might affect dominant species relying on chemical recruitment, more than subordinate ant species, less dependent on chemical communication and less sensitive to high temperatures.

Make love not war: when should less competitive males choose low-quality but defendable females?

Male choosiness for mates is an underexplored mechanism of sexual selection. A few theoretical studies suggest that males may exhibit--but only under rare circumstances--a reversed male mate choice (RMMC; i.e., highly competitive males focus on the most fecund females, while the low-quality males exclusively pair with less fecund mates to avoid being outcompeted by stronger rivals). Here we propose a new model to explore RMMC by relaxing some of the restrictive assumptions of the previous models and by considering an extended range of factors known to alter the strength of sexual selection (males' investment in reproduction, difference of quality between females, operational sex ratio). Unexpectedly, we found that males exhibited a reversed mate choice under a wide range of circumstances. RMMC mostly occurs when the female encounter rate is high and males devote much of their time to breeding. This condition-dependent strategy occurs even if there is no risk of injury during the male-male contest or when the difference in quality between females is small. RMMC should thus be a widespread yet underestimated component of sexual selection and should largely contribute to the assortative pairing patterns observed in numerous taxa.

What matters in the associative learning of visual cues in foraging parasitoid wasps: colour or brightness?

Visual cues are known to be used by numerous animal taxa to gather information on quality and localisation of resources. Because environmental lighting can interfere with the spectral features of visual cues, the specific characteristics of the colour signals that promote forager decision and learning are still not known in the majority of insects (excepted in bees). We analysed the effect of previous experience on the use of visual information by the wasp Venturia canescens, a parasitoid of pyralidae, in the context of host searching. These parasitoids search for hosts concealed in several fruit species, so visual cues from the host microhabitat could play a key role in host finding. We also investigated the type of visual cues on which wasps based their decision. We tested whether wasps are able to associate an achromatic cue (brightness) or a chromatic one (hue, i.e. dominant wavelength and/or chroma) with the presence of hosts. Our results show that in the context of host foraging, chromatic cues are more reliable than brightness in achieving the associative learning process. Therefore, understanding the behavioural ecology of foraging should make use of the knowledge about the visual information used.

Dominance-discovery and discovery-exploitation trade-offs promote diversity in ant communities.

In ant communities, species coexist by using different foraging strategies. We developed an adaptive dynamics model to gain a better understanding of the factors that promote the emergence and maintenance of strategy diversity. We analysed the consequences of both interspecific competition and resource distribution for the evolutionary dynamics of social foraging in ants. The evolution of social foraging behaviour was represented using a stochastic mutation-selection process involving interactions among colonies. In our theoretical community, ant colonies inhabit an environment where resources are limited, and only one resource type is present. Colony interactions depend on colony-specific foraging strategies (defined as the degree of collective foraging), resource distribution patterns, and the degree of competition asymmetry. At the ecological timescale, we have created a model of foraging processes that reflects trade-offs between resource discovery and resource exploitation and between resource discovery and ant behavioural dominance. At the evolutionary timescale, we have identified the conditions of competition and resource distribution that can lead to the emergence and coexistence of both collective and individual foraging strategies. We suggest that asymmetric competition is an essential component in the emergence of diverse foraging strategies in a sympatric ant community.

Adaptations to different habitats in sexual and asexual populations of parasitoid wasps: a meta-analysis.

BACKGROUND: Coexistence of sexual and asexual populations remains a key question in evolutionary ecology. We address the question how an asexual and a sexual form of the parasitoid Venturia canescens can coexist in southern Europe. We test the hypothesis that both forms are adapted to different habitats within their area of distribution. Sexuals inhabit natural environments that are highly unpredictable, and where density of wasps and their hosts is low and patchily distributed. Asexuals instead are common in anthropic environments (e.g., grain stores) where host outbreaks offer periods when egg-load is the main constraint on reproductive output. METHODS: We present a meta-analysis of known adaptations to these habitats. Differences in behavior, physiology and life-history traits between sexual and asexual wasps were standardized in term of effect size (Cohen's d value; Cohen, 1988). RESULTS: Seeking consilience from the differences between multiple traits, we found that sexuals invest more in longevity at the expense of egg-load, are more mobile, and display higher plasticity in response to thermal variability than asexual counterparts. DISCUSSION: Thus, each form has consistent multiple adaptations to the ecological circumstances in the contrasting environments.

Predator Migration Decisions, the Ideal Free Distribution, and Predator-Prey Dynamics.

The aim of the present work is to analyze the influence of optimal predator emigration decisions that can lead to the ideal free distribution (IFD) on the stability of predator-prey systems. The assumption of optimal decisions is then relaxed to analyze the possible influence of different degrees of deviation from the IFD. The first migration rule we analyze is based on the marginal-value theorem and assumes perfect knowledge of capture rate in the patch of residence and in the environment as a whole. When migration rates are high, this rule leads the predator population to the IFD. The results suggest that under these conditions predator migration plays no major role in the stability of the system. This is so because the systems naturally merge into a single patch. This result is independent of the particular functional response used. The other two rules we analyze take into account lower migration rates, the limitations in making optimal decisions by predators, and the possible constraints in the assessment of intake rate in the different patches. The results suggest that the processes that hinder the convergence of the populations to the IFD might make a major contribution to the stability of the system.

Energy dynamics in a parasitoid foraging in the wild.

Although parasitoids are used widely as a biological models for understanding the evolution of animal behaviour, most studies have been constrained to the laboratory. The dearth of field studies has been compounded by the almost complete ignorance of the physiological parameters involved in foraging and dispersal, in particular of the energetic constraints imposed by resource limitation. We estimated the dynamics of carbohydrates and lipids reserves of Venturia canescens (Gravenhorst) females by releasing individuals of known nutritional status in a natural environment and recapturing them using host-containing traps. The recapture rate was around 30%. These results were compared with the reserves of caged animals kept under different experimental conditions (freshly emerged, starved to death, fed ad libitum and partially starved). Wild animals were also sampled in order to estimate the resource levels of the local population. The results show that: (i) wasps are able to maintain a nearly constant level of energy over an extended foraging period; (ii) V. canescens takes sugars in the field; and (iii) the lipid reserves accumulated during the larval life may be limiting as lipogenesis does not take place in adults even under conditions of high sugar availability. These results demonstrate that wasps can forage for hosts and food and disperse in this habitat for hours and days without running into a severe risk of energy limitation.

Density dependence and regulation in the systemNezara viridula (L.) (Hemiptera: Pentatomidae), host andTrichopoda giacomellii (Blanchard) (Diptera: Tachinidae), parasitoid.

The system formed byN. viridula and its almost specific parasitoidT. giacomellii has been studied in the Buenos Aires province, Argentina for 26 generations of the former and 20 generations of the latter. An intensive study over 9 host generations and key factor analysis made possible the identification and quantification of different mortality factors. The main factors determining the fluctuations in total generation mortality were: egg mortality due mainly to parasitism byT. basalis; nymphal predation on 1st, 2nd and 3rd instar nymphs and migration; and adult mortality and reduction on potential natality due to parasitism byT. giacomellii. Adult mortality and reduction of potential natality, due to parasitism byT. giacomellii, was density dependent. This factor is capable of maintaining the simulations generated by a simple model based on key factor analysis, within the range of observed densities. These results suggest that parasitism byT. giacomellii can regulate the population, even in a small area of 450 m2.

Does a deletion in a virus-like particle protein have pleiotropic effects on the reproductive biology of a parasitoid wasp?

Endoparasitoid insects have evolved mechanisms to counteract host immune defences. At oviposition, the endoparasitoid Venturia canescens injects virus-like particles (VLPs) together with the egg that interfere with the immune system of the host. These prevent encapsulation of the parasitoid egg. It has been shown that the gene coding for one of the VLP proteins exists in two variants (called Repeat-Plus (RP) and Repeat-Minus (RM)). Previous observations suggested that these variants induce pleiotropic effects on the reproductive biology of the female resulting, in an impeded transfer of eggs from the ovarioles to the oviducts in RM females, and in alterations in the egg laying sequence. We show that RM females from another geographical locality do not exhibit any phenotypic alteration of the reproductive biology. By showing a lack of association between VLP alleles and the reproductive phenotypic characteristics in a given strain, our results do not support the hypothesis of pleiotropic effects. Conversely, the results suggest that different genes code for the VLPs and for the reproductive biology characteristics. Different phenomena such as linkage, the action of pathogens, etc. could explain the association between characters that is observed in some strains.

A list of and some comments about the trail pheromones of ants.

Ants use many different chemical compounds to communicate with their nestmates. Foraging success depends on how efficiently ants communicate the presence of food and thus recruit workers to exploit the food resource. Trail pheromones, produced by different exocrine glands, are a key part of ant foraging strategies. By combing through the literature, we compiled a list of the identity and glandular origin of the chemical compounds found in the trail pheromones of 75 different ant species. Of the 168 compounds identified, more than 40% are amines. In the subfamily Myrmicinae, trail pheromones are mostly produced in the venom gland, while in the subfamily Formicinae, they come from the rectal gland.

An evolutionary dynamics model adapted to eusocial insects.

This study aims to better understand the evolutionary processes allowing species coexistence in eusocial insect communities. We develop a mathematical model that applies adaptive dynamics theory to the evolutionary dynamics of eusocial insects, focusing on the colony as the unit of selection. The model links long-term evolutionary processes to ecological interactions among colonies and seasonal worker production within the colony. Colony population dynamics is defined by both worker production and colony reproduction. Random mutations occur in strategies, and mutant colonies enter the community. The interactions of colonies at the ecological timescale drive the evolution of strategies at the evolutionary timescale by natural selection. This model is used to study two specific traits in ants: worker body size and the degree of collective foraging. For both traits, trade-offs in competitive ability and other fitness components allows to determine conditions in which selection becomes disruptive. Our results illustrate that asymmetric competition underpins diversity in ant communities.

Does kin recognition and sib-mating avoidance limit the risk of genetic incompatibility in a parasitic wasp?

BACKGROUND: When some combinations of maternal and paternal alleles have a detrimental effect on offspring fitness, females should be able to choose mates on the basis of their genetic compatibility. In numerous Hymenoptera, the sex of an individual depends of the allelic combination at a specific locus (single-locus Complementary Sex Determination), and in most of these species individuals that are homozygous at this sexual locus develop into diploid males with zero fitness. METHODS AND FINDINGS: In this paper, we tested the hypothesis of genetic incompatibility avoidance by investigating sib-mating avoidance in the solitary wasp parasitoid, Venturia canescens. In the context of mate choice we show, for the first time in a non-social hymenopteran species, that females can avoid mating with their brothers through kin recognition. In "no-choice" tests, the probability a female will mate with an unrelated male is twice as high as the chance of her mating with her brothers. In contrast, in choice tests in small test arenas, no kin discrimination effect was observed. Further experiments with male extracts demonstrate that chemical cues emanating from related males influence the acceptance rate of unrelated males. CONCLUSIONS: Our results are compatible with the genetic incompatibility hypothesis. They suggest that the female wasps recognize sibs on the basis of a chemical signature carried or emitted by males possibly using a "self-referent phenotype matching" mechanism.

Flight variability in the woodwasp Sirex noctilio (Hymenoptera: Siricidae): an analysis of flight data using wavelets.

We describe flight variability in the woodwasp Sirex noctilio Fabricius, 1793 (Hymenoptera: Siricidae) by studying tethered females in a flight mill device and analyzing output data by a time series methodology. Twenty-eight wasps were flown during 24 h-long periods, under controlled temperature and lighting conditions. The maximum distance recorded was 49 km, and mean velocity was 0.37 m s(-1). All wasps lost weight during flight (mean weight loss of 10.0% of initial body mass). By using a wavelets analysis on the flight mill time series output, we identified three distinct flight patterns: regular (long acceleration-deceleration spells), periodic (alternation of acceleration-deceleration spells without resting) and pulsating (resting spells interrupted by bursts of flight activity). The first two flight patterns are indistinguishable using traditional flight mill data analysis. Flight patterns for each individual were significantly dependent on wasp body mass, suggesting a relationship with the resources used in flight and their availability. Large females flew sequentially through a regular-periodic-pulsating sequence but medium sized wasps flew mostly with periodic and pulsating patterns. The smallest wasps flew only in a pulsating pattern, being incapable of long, sustained flight. Variability in size and behavior can have significant consequences on population dynamics by determining local and regional dispersal. An important outcome of our work is the introduction of wavelet analysis to study tethered flight data series for the first time. This methodology allowed us to uncover and statistically test individual variability in insect flight characteristics.

Foraging and associative learning of visual signals in a parasitic wasp.

To cope with environmental variability, animals should gather and use information to reduce uncertainty. In insect parasitoids, associative learning has been widely documented in the context of host foraging. However, despite its potential adaptive value, the insect food searching strategy and cues used to search are poorly understood. In this study, we examined the ability of hymenopteran Venturia canescens females to associate food to a visual cue. To broaden the scope of our results, experiments were performed with both arrhenotokous (sexual) and thelytokous (asexual) individuals. The wasps showed innate attraction for yellow and orange stimuli when presented versus blue stimuli. When trained to associate a food reward with one of the attractive colours (orange), they significantly moved from a distance towards the colour previously associated with food. The choice of the innately preferred colour (yellow) was not modified by associative learning. In the context of food foraging, this study is the first to show associative learning using visual stimuli in a parasitoid and active choice of this colour. This ability gives new insights concerning potential food sources for V. canescens in the field, since flowers are sugar sources, which emit colour signals.

The influence of temperature and host availability on the host exploitation strategies of sexual and asexual parasitic wasps of the same species.

In the hymenopteran parasitoid Venturia canescens, asexual (obligate thelytoky not induced by Wolbachia bacteria) and sexual (arrhenotokous) wasps coexist in field conditions despite the demographic cost incurred due to the production of males by sexual females. Arrhenotoky predominates in field conditions, whereas populations in indoor conditions (mills, granaries) are exclusively thelytokous. These differences in the relative abundance of the two modes of reproduction between environments suggest that the individuals of each reproductive mode may have developed strategies adapted to the conditions prevailing in each kind of habitat. The two environments contrast in temperature variability and in the spatial heterogeneity of host availability. In this study, we considered the combined effect of temperature and host availability on host patch exploitation by thelytokous and arrhenotokous V. canescens. As expected, arrhenotokous females were more sensitive to temperature changes. If the temperature decreased before foraging, they remained longer and exploited patches more thoroughly. This is consistent with the expected behaviour of parasitoids in response to signs of unfavourable conditions that entail increasing risk of time limitation or a reduced probability of attaining further patches. Both arrhenotokous and thelytokous females increased patch exploitation with host availability. However, unexpectedly, we found no difference in the way the two types of wasp responded to differences in host availability. Differences in the strategies adopted under different environmental conditions may indicate divergence of niche-specific life history traits between the two modes of reproduction. Niche displacement may partly account for the coexistence of these two modes of reproduction at a geographical scale.