Influence of Inbreeding on Female Mate Choice in Two Species of
Many organisms have been reported to choose their mates in order to increase the heterozygosity of their offspring by avoiding mating with relatives or homozygous individuals. Most previous studies using have used artificial chromosomes or extreme inbreeding treatments, situations unlikely to be matched in nature. Additionally, few studies have examined the interaction between female inbreeding status and her choice of mate. Using females and males from populations that had experienced either random mating or one generation of sib-sib inbreeding, we measured the preferences of females for males. Our results indicate that outbred males were chosen more often than inbred males and that this preference may be more pronounced in outbred females than in inbred ones.
Differing Host Exploitation Efficiencies in Two Hyperparasitoids: When is a 'Match Made in Heaven'?
Host exploitation behavior in two hyperparasitoids, Lysibia nana and Gelis agilis, was compared in single cocoon clusters of their primary parasitoid host, Cotesia glomerata. L. nana reproduces sexually, is fully winged, does not host-feed and matures eggs quite rapidly after eclosion, whereas G. agilis possesses opposite traits. Cohorts of individual hyperparasitoid females of differing age and physiological state were given access to single cocoon clusters of C. glomerata that also varied in age. These results reveal that the reproductive biology of L. nana is well matched to exploit cocoon broods in C. glomerata, suggesting strong a co-evolutionary history with this host. By contrast, G. agilis is much less efficient at exploiting host cocoons and is probably a generalist species that attacks other hosts in nature.
Parasite Lost: Chemical and Visual Cues Used by Pseudacteon in Search of Azteca instabilis
An undescribed species of phorid fly (genus: Pseudacteon) parasitizes the ant Azteca instabilis F Smith, by first locating these ants through the use of both chemical and visual cues. Experiments were performed in Chiapas, Mexico to examine a) the anatomical source of phorid attractants, b) the specific chemicals produced that attract phorids, and c) the nature of the visual cues used by phorids to locate the ants. We determined that phorid-attracting chemicals were present within the dorsal section of the abdomen, the location of the pygidial gland. Further experiments indicate that a pygidial gland compound, 1-acetyl-2-methylcyclopentane, is at least partially responsible for attracting phorid flies to their host. Finally, although visual cues such as movement were important for host location, size and color of objects did not influence the frequency with which phorids attacked moving targets.
Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition
Collective decision making in the social insects often proceeds via feedback cycles based on positive signaling. Negative signals have, however, been found in a few contexts in which costs exist for paying attention to no longer useful information. Here we incorporate new research on the specificity and context of the negative stop signal into an agent based model of honey bee foraging to explore the adaptive basis of negative signaling in the dance language. Our work suggests that the stop signal, by acting as a counterbalance to the waggle dance, allows colonies to rapidly shut down attacks on other colonies. This could be a key adaptation, as the costs of attacking a colony strong enough to defend itself are significant. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1007/s10905-010-9229-5) contains supplementary material, which is available to authorized users.
Effects of Intraspecific Competition and Host-Parasitoid Developmental Timing on Foraging Behaviour of a Parasitoid Wasp
In a context where hosts are distributed in patches and susceptible to parasitism for a limited time, female parasitoids foraging for hosts might experience intraspecific competition. We investigated the effects of host and parasitoid developmental stage and intraspecific competition among foraging females on host-searching behaviour in the parasitoid wasp We found that females have a pre-reproductive adult stage during which their eggs are not mature yet and they forage very little for hosts. The wasps foraged for hosts more once they were mature. Behavioural experiments showed that wasps' foraging activity also increased as host eggs aged and became susceptible to parasitism, and as competition among foraging wasps increased.
Triggers of the Postural Display of Courtship in Flies
courtship shows some flexibility and courting males sometimes perform an elaborate postural display in addition to the standard courtship behaviours shared by most species. This postural display includes the acrobatic contortion and tremulation of their abdomen, accompanied by the generation of substrate-borne vibrations, and they proffer a nutritional droplet to the female. Here, we use courtship and choice assays to ask what triggers this display and what advantages males may gain from it during courtship. In pair assays, we found no differences in the courtship duration and copulation success between displaying and non-displaying males. In trio assays, however, the female always mated with the male who performed the display. To investigate what promotes the male display, we varied the level of receptivity of the female and studied the impact of a second male. We found that rejection by the female does not induce the male to display, contrary to what was previously suggested. We present evidence that the male display is in fact promoted by the presence of an attentive and sexually receptive female and the absence of male competition, with the greatest exhibition rate obtained if the courted female is starved. These findings provide valuable information about the social ecology of flies, and how internal and external cues influence sexual behaviours and mate choice.
Behavioral Reversion and Dark-Light Choice Behavior in Workers of the Red Wood Ant
Social insect workers usually start adult life from intranidal tasks and then switch to extranidal activities, but this process may be reversed: foragers may switch again to intranidal brood care. The transition forager - reverted nurse is known as the behavioral reversion. Ant foragers are known to avoid illuminated zones less strongly than intranidal workers, but illumination responses of reverted nurses were so far never investigated. We compared dark-light choice behavior of three classes of workers of the red wood ant : nurses, foragers and reverted nurses. Sets of ten ants belonging to the same class were tested in "double nests" made of two interconnected test tubes, one kept in darkness and the other exposed to light. The number of ants present in the illuminated zone of each nest (n) was recorded on 10 sample points at 30 min intervals. The values of n were lower in nurses than in foragers and reverted nurses and decreased as a function of time in all three groups. Nurses differed from foragers with respect to the dynamics of dark-light choice behavior, but reverted nurses did not differ in that respect either from nurses, or from foragers. Reverted nurses and foragers did not differ significantly from each other with respect to the overall level of avoidance of illuminated zone, nor with respect to the dynamics of dark-light choice behavior. This implies that behavioral reversion is not accompanied by the return of illumination responses of workers of to the state characteristic for nurses.
Primacy of Human Odors Over Visual and Heat Cues in Inducing Landing in Female Mosquitoes
Although human skin odor is thought to be the cue that anthropophilic mosquitoes use to discriminate us from other potential hosts, the precise details of how they use skin odor to find and land on a human is unclear. We found that land on a source of skin odor without a co-located visual cue. By collecting human odor on glass beads and using identical glass beads to visually conceal skin odor and heat cues, we were able to study mosquito landing on skin odor, heat, and visual cues separately. Landing is necessary for blood feeding which is a required behavior for the life cycle as well as the behavior responsible for the epidemiological impact of mosquitoes. Therefore, we consider it to be the diagnostic measure of the importance of a host cue. In two-choice tests, a skin odor source had the highest valence for landing, followed by a combination of heat and a visual cue, and finally heat and visual cues presented separately. We also measured the durations of the landings, though no significant differences were found.
The Elaborate Postural Display of Courting Flies Produces Substrate-Borne Vibratory Signals
Sexual selection has led to the evolution of extraordinary and elaborate male courtship behaviors across taxa, including mammals and birds, as well as some species of flies. flies perform complex courtship behaviors found in most species, which consist of visual, air-borne, gustatory and olfactory cues. In addition, courting males also perform an elaborate postural display that is not found in most other species. This postural display includes an upwards contortion of their abdomen, specialized movements of the head and forelegs, raising both wings into a "wing-posture" and, most remarkably, the males proffer the female a regurgitated droplet. Here, we use high-resolution imaging, laser vibrometry and air-borne acoustic recordings to analyse this postural display to ask which signals may promote copulation. Surprisingly, we find that no air-borne signals are generated during the display. We show, however, that the abdomen tremulates to generate substrate-borne vibratory signals, which correlate with the female's immobility before she feeds onto the droplet and accepts copulation.
Floral Sonication is an Innate Behaviour in Bumblebees that can be Fine-Tuned with Experience in Manipulating Flowers
Bumblebees demonstrate an extensive capacity for learning complex motor skills to maximise exploitation of floral rewards. This ability is well studied in nectar collection but its role in pollen foraging is less well understood. Floral sonication is used by bees to extract pollen from some plant species with anthers which must be vibrated (buzzed) to release pollen. Pollen removal is determined by sonication characteristics including frequency and amplitude, and thus the ability to optimise sonication should allow bees to maximise the pollen collection. We investigated the ability of the buff-tailed bumblebee () to modify the frequency and amplitude of their buzzes with increasing experience manipulating flowers of the buzz-pollinated plant . We analysed flight and feeding vibrations generated by naïve workers across feeding bouts. Feeding buzzes were of a higher frequency and a lower amplitude than flight buzzes. Both flight and feeding buzzes had reduced amplitudes with increasing number of foraging trips. However, the frequency of their feeding buzzes was reduced significantly more than their flight buzzes as bumblebee workers gained experience manipulating flowers. These results suggest that bumblebees are able to modify the characteristics of their buzzes with experience manipulating buzz-pollinated flowers. We discuss our findings in the context of bumblebee learning, and the current understanding of the optimal sonication characteristics for releasing pollen in buzz-pollinated species. Our results present a tantalising insight into the potential role of learning in floral sonication, paving the way for future research in this area.
The Effect of Wind Exposure on the Web Characteristics of a Tetragnathid Orb Spider
Studies on spiders in their natural habitats are necessary for determining the full range of plasticity in their web-building behaviour. Plasticity in web design is hypothesised to be important for spiders building in habitats where environmental conditions cause considerable web damage. Here we compared web characteristics of the orb spider (Araneae, Tetragnathidae) in two different forest habitats differing in their wind exposure. We found a notable lack of differences in web geometry, orientation and inclination between webs built along an exposed forest edge and those built inside the forest, despite marked differences in wind speed. This suggests that did not exhibit web-building plasticity in response to wind in the field, contrasting with the findings of laboratory studies on other species of orb spiders. Instead, differences in prey capture and wind damage trade-offs between habitats may provide an explanation for our results, indicating that different species employ different strategies to cope with environmental constraints.
Attraction of the Larval Parasitoid (Hymenoptera: Pteromalidae) to Feces Volatiles from the Adult Weevil Host
The behavioral response of the larval parasitoid (Hymenoptera: Pteromalidae) to volatile compounds derived from its weevil hosts was investigated in two-choice bioassays. Odor source candidates were the larval and adult stages of weevils, clover flowers, and feces from adult weevils. Despite being a larval parasitoid, the odor of weevil larvae isolated from the clover flowers was not attractive to female parasitoids. Surprisingly, females were instead attracted by the odor from the feces of adult weevils. The female parasitoids were similarly attracted to the feces produced by the two main hosts, the red clover weevil () and the white clover weevil () Chemical analysis of the volatile composition of feces produced by the two hosts revealed qualitatively similar odor profiles, correlating with the observed attraction by the parasitoid towards both odor sources. Some of the identified volatile compounds are commonly present in clover plant headspace fractions and may function as a kairomone to facilitate orientation by to -infested clover flowers. Larval and adult weevils were not attractive for parasitoid females, whereas, for the white clover weevil-plant association, infested flowers were highly attractive. These data show the use by the clover weevil parasitoid of an alternative source of olfactory information for locating its host.
Investigating the Foraging, Guarding and Drifting Behaviors of Commercial
Social insects have high levels of cooperation and division of labor. In bumble bees this is partly size-based, with larger bees performing tasks outside the nest and smaller bees remaining inside, although bumble bees still display considerable behavioral plasticity. The level of specialization in tasks outside the colony, including foraging, guarding and drifting (entering a foreign colony), is currently unknown for bumble bees. This study aimed to assess division of labor between outside tasks and the degree of specialization in foraging, guarding, and switching colonies in commercially reared bumble bees placed in the field. Nine factory-bought colonies were placed on three farms in Sussex, UK, between June and August 2015. Forty workers from each colony were radio-tagged and a reader on the colony entrance recorded the date, time and bee ID as they passed. The length and frequency of foraging trips and guarding behavior were calculated, and drifting recorded. The mean (±SD) length of foraging trips was 45 ± 36 min, and the mean number of foraging trips per day was 7.75 ± 7.71. Low levels of specialization in guarding or foraging behavior were found; however, some bees appeared to guard more frequently than others, and twenty bees were categorized as guards. Five bees appeared to exhibit repeated "stealing" behavior, which may have been a specialist task. The division of labor between tasks was not size-based. It is concluded that commercial bumble bees are flexible in performing outside nest tasks and may have diverse foraging strategies including intra-specific nest robbing.
Functional and Behavioral Responses of the Natural Enemy to , at Different Temperatures
is the dominant predator of pear sucker () in the UK. migrates into orchards in spring or is introduced as a biocontrol agent, reaching peak population levels in July-August, contributing to effective control of summer pear sucker populations. However, due to temperature dependent development and metabolism there are concerns that populations or feeding rates may increase due to changing climatic conditions. Thus, how responds to temperature, impacts its ability as a biocontrol agent. Functional response assays, monitoring attack rate and handling time of and behavioral assays, using Ethovision tracking software occurred, to assess the impact of temperature on predation. Experiments were conducted at current and future July-August mean temperatures, predicted using RCP4.5 and RCP8.5 (medium and high, representative concentration pathway) emissions scenarios, using 2018 UK Climate Projections (UKCP18). All treatments demonstrated a Type II functional response, with female anthocorids demonstrating shorter handling times and higher attack rates than males. Males showed longer prey handling times at 18 °C compared to 23 °C and more time was spent active at lower temperatures for both sexes. Females did not show significant differences in attack rate or handling time in response to temperature. Overall prey consumption was also not significantly affected by temperature for either sex. This study suggests that anthocorids are likely to remain effective natural enemies under future predicted temperatures, due to non-significant differences in prey consumption.
Increased Risk Proneness or Social Withdrawal? The Effects of Shortened Life Expectancy on the Expression of Rescue Behavior in Workers of the ant (Hymenoptera: Formicidae)
In social insects behavioral consequences of shortened life expectancy include, among others, increased risk proneness and social withdrawal. We investigated the impact of experimental shortening of life expectancy of foragers of the ant achieved by their exposure to carbon dioxide on the expression of rescue behavior, risky pro-social behavior, tested by means of two bioassays during which a single worker (rescuer) was confronted with a nestmate (victim) attacked by a predator (antlion larva capture bioassay) or immobilized by an artificial snare (entrapment bioassay). Efficacy of carbon dioxide poisoning in shortening life expectancy was confirmed by the analysis of ant mortality. Rescue behavior observed during behavioral tests involved digging around the victim, transport of the sand covering the victim, pulling the limbs/antennae/mandibles of the victim, direct attack on the antlion (in antlion larva capture tests), and snare biting (in entrapment tests). The rate of occurrence of rescue behavior was lower in ants with shortened life expectancy, but that effect was significant only in the case of the entrapment bioassay. Similarly, only in the case of the entrapment bioassay ants with shortened life expectancy displayed rescue behavior after a longer latency and devoted less time to that behavior than ants from the control groups. Our results demonstrated that in ant workers shortened life expectancy may lead to reduced propensity for rescue behavior, most probably as an element of the social withdrawal syndrome that had already been described in several studies on behavior of moribund ants and honeybees.
Aphids Playing Possum - Defensive or Mutualistic Response?
Paper presents the phenomenon of thanatosis or death-feigning in selected aphids species. This specific reaction was predominantly analysed on the example of aphid subfamily Lachninae. Individuals of this group were used in experiments, during which a thanatotic response was induced with various results. The response differed from prolonged thanatosis, lasting for several minutes (), through shrinking behaviour (e. g. in ) to non-responsive species such as (i) . The large interspecies variation of observed responses can be linked to other defensive mechanisms existing in the studied species, as well as to their mutualistic relationship with ants. The behaviour of shrinking is hypothesized to be the mutualistic response, developed from thanatosis, and being adapted to transportation by ant workers.
Behavioral Plasticity in Probing by (Hemiptera, Liviidae): Ingestion from Phloem Versus Xylem is Influenced by Leaf Age and Surface
is a major pest of citrus because it transmits Liberibacter asiaticus, a phloem-limited bacterium that putatively causes Huanglongbing (HLB). The disease moves slowly through a tree, and the vector facilitates further within-tree movement via transmission of the pathogen. However, this only happens when stylets contact the phloem, to inoculate bacteria during phloem salivation and acquire bacteria during phloem sap ingestion. Behavioral changes in associated with different plant parts would affect how long it takes to reach phloem and how long the psyllids stays in phloem to ingest, thereby influencing the risk of disease spread. feeding was recorded on the abaxial and adaxial surfaces of mature and immature citrus leaves. Adults in the field can be found on these surfaces at all times of year. On abaxial surface of immature leaves, phloem salivation would occur after 11 h on average, but rarely as soon as 0.56 h. The corresponding values on mature leaves were 16 and 2.7. In general, psyllids spent more time ingesting phloem sap on immature leaves than on mature leaves. Psyllids on abaxial surfaces spent more time ingesting from phloem, though the strength of this effect was less than for immature versus mature leaves. In contrast, xylem ingestion increased on mature leaves compared with young. The biological differences that could produce this outcome are discussed. The results discussed herein are of relevance to further studies on the efficacy of an insecticide to act quickly enough to prevent pathogen transmission.
Aversive Learning in the Praying Mantis (), a Sit and Wait Predator
Animals learn to associate sensory cues with the palatability of food in order to avoid bitterness in food (a common sign of toxicity). Associations are important for active foraging predators to avoid unpalatable prey and to invest energy in searching for palatable prey only. However, it has been suggested that sit-and-wait predators might rely on the opportunity that palatable prey approach them by chance: the most efficient strategy could be to catch every available prey and then decide whether to ingest them or not. In the present study, we investigated avoidance learning in a sit-and-wait predator, the praying mantis (). To examine the effects of conspicuousness and novelty of prey on avoidance learning, we used three different prey species: mealworms (novel prey), honeybees (novel prey with conspicuous signals) and crickets (familiar prey). We sequentially presented the prey species in pairs and made one of them artificially bitter. In the absence of bitterness, the mantises consumed bees and crickets more frequently than mealworms. When the prey were made bitter, the mantises still continued to attack bitter crickets as expected. However, they reduced their attacks on bitter mealworms more than on bitter bees. This contrasts with the fact that conspicuous signals (e.g. coloration in bees) facilitate avoidance learning in active foraging predators. Surprisingly, we found that the bitter bees were totally rejected after an attack whereas bitter mealworms were partially eaten (~35%). Our results highlight the fact that the mantises might maintain a selection pressure on bees, and perhaps on aposematic species in general.
Behavior of the Avian Parasite (Diptera: Muscidae) in and Near Host Nests in the Galapagos Islands
The Avian Vampire Fly, , has invaded the Galapagos Islands, where it causes high mortality of endemic and native landbird species, including most species of Darwin's finches. Control methods are under development, but key information is missing about the reproductive biology of and the behavior of flies in and near nests of their hosts. We used external and internal nest cameras to record the behavior of adults within and outside nests of the Galapagos Flycatcher, , throughout all stages of the nesting cycle. These recordings showed that visited flycatcher nests throughout the day with higher fly activity during the nestling phase during vespertine hours. The observations also revealed that multiple individuals can visit nests concurrently, and that there are some interactions among these flies within the nest. Fly visitation to nests occurred significantly more often while parent birds were away from the nest than in the nest, and this timing appears to be a strategy to avoid predation by parent birds. We report fly mating behavior outside the nest but not in the nest cavity. We discuss the relevance of these findings for the adaptive forces shaping life history strategies as well as rearing and control measures.
Predation Cues in Solitary bee Nests
Predation at the nesting site can significantly affect solitary bees' reproductive success. We tested female red mason bees' ( L.) acceptance of potential nesting sites, some of which were marked with cues coming from predated conspecifics (crushed bees) or from a predator itself (rodent excreta). In our experiment, females did not avoid nests marked with either of the two predator cues. We suggest that bee females do not recognize these two cues as risky. Alternatively, costs of abandoning natal aggregation might be too high compared with any perceived predation risk of staying. Moreover, the presence of crushed bees can provide positive information about the presence of conspecifics and, possibly, information about a nesting aggregation that may be preferred by bees when choosing a nesting site.
Traits Underlying Experimentally Evolved Dispersal Behavior in
Dispersal is an important behavior in many animals, with profound effects on individual fitness and the evolutionary trajectories of populations. This is especially true within taxa with particular life-history strategies, for example those that exploit ephemeral habitat. Further, dispersal is commonly seen to be part of behavioral syndromes - suites of traits that covary across behavioral contexts. The red flour beetle, (Coleoptera, Tenebrionidae), is a major post-harvest crop pest responsible for large losses through the infestation of stored grain. In this system dispersal is known to have a strong genetic basis and differential artificial selection on dispersal traits produces strong phenotypic divergence. However, it is unknown which traits are able to rapidly evolve to produce these results, or which behavioral components underlie differences in dispersal. Using replicate lines of previously selected for divergent dispersal behavior, we test for correlated activity and movement patterns, morphology and substrate surface use. We find robustly repeatable associations between the dispersal phenotype and higher activity, straighter paths, larger body size (but not relative leg length) and increased tendency to remain at the surface of fodder. Together our results suggest that dispersal is part of a syndrome of traits in , and must be treated as such when considering the evolution of dispersal in this system, and in attempting to predict and control its spread.
