Neural control of homologous behaviour patterns in two blaberid cockroaches

Activity patterns of motoneurones which innervate spiracular muscles in two blaberid cockroaches, Blaberus discoidalis and Gromphadorhina portentosa, have been monitored during two homologous behaviour patterns: respiratory and non-respiratory tracheal ventilation. Based upon the activity of spiracular motoneurones during these two activities, the abdominal spiracles have been divided into three functional groups: vestigial, respiratory and non-respiratory. In Blaberus discoidalis spiracle 3 is vestigial, spiracles 6, 7, 8 and 10 are respiratory, and spiracles 4, 5 and 9 are non-respiratory. In Gromphadorhina portentosa spiracles 3 and 10 are vestigial, spiracle 4 is non-respiratory and spiracles 5–9 are respiratory.Respiratory spiracles in both species are characterized by activity patterns of their motoneurones during respiratory tracheal ventilation: low frequency firing at irregular intervals during the respiratory pause and a higher frequency burst synchronous with the expiratory abdominal compression. Non-respiratory spiracles are characterized by complete inactivity of their opener motoneurones during respiratory tracheal ventilation. These motoneurones are activated by mechanical stimulation in both species, which simultaneously suppresses activity in respiratory opener motoneurones. In Blaberus discoidalis, there are no differences between activity patterns of respiratory and non-respiratory closer motoneurones. In Gromphadorhina portentosa, not only do respiratory and non-respiratory closer motoneurones have different activity patterns, but the activity pattern of respiratory closer motoneurones is different during respiratory and non-respiratory tracheal ventilation. The functional implications of these several spiracular motoneurone activity patterns are discussed.     

Questing activity in bed bug populations: male and female responses to host signals

A large‐arena bioassay is used to examine sex differences in spatiotemporal patterns of bed bug Cimex lectularius L. behavioural responses to either a human host or CO₂ gas. After release in the centre of the arena, 90% of newly‐fed bed bugs move to hiding places in the corners within 24 h. They require 3 days to settle down completely in the arena, with generally low activity levels and the absence of responses to human stimuli for 5 days. After 8–9 days, persistent responses can be recorded. Sex differences are observed, in which females are more active during establishment, respond faster after feeding, expose themselves more than males during the daytime, and respond more strongly to the host signal. The number of bed bugs that rest in harbourages is found to vary significantly according to light setting and sex. Both sexes stay inside harbourages more in daylight compared with night, and males hide more than females during the daytime but not during the night. The spatial distribution of the bed bugs is also found to change with the presence of CO₂, and peak aggregation around the odour source is observed after 24 min. Both male and female bed bugs move from hiding places or the border of the arena toward the centre where CO₂ is released. Peak responses are always highest during the night. Bed bug behaviour and behaviour‐regulating features are discussed in the context of control methods.     

Female House Mice do not Differ in Their Exploratory Behaviour from Males

Dispersal is an important ecological trait affecting genetic variation and dynamics of populations. Hence, the exploratory behaviour prior to actual dispersal may be crucial for potentially dispersing individuals. In mammals, females are traditionally seen as the more philopatric sex and dispersal as male‐biased behaviour, and so behavioural strategies related to the exploration of novel resources should be differentially expressed in males and females. In addition, due to sexual selection exploratory strategies may be expected to vary according to females’ reproductive phase. We employed a standard open‐field test as an approximation of the first phase of dispersal, using adult house mice representing two subspecies, M. m. musculus and M. m. domesticus. We tested the prediction that exploration of neutral area varies in females during different phases of the oestrus cycle and is different between both sexes and subspecies. We expected to find higher exploration in males, as the more dispersing sex and less pronounced subspecies‐specific differences in females than in males. We found no significant effect of the oestrous phase on any of the parameters of the exploratory behaviour measured. Sexual dimorphism was found only in latency to enter the arena in M. m. domesticus where females hesitated longer to enter a new area than males. Significant subspecies‐specific differences were found in three of four tested exploration parameters, so we conclude that females of both subspecies follow similar strategies to those displayed by males. Musculus mice show shorter latency to enter a new area, but once inside, domesticus mice explore the arena significantly longer, with less frequent retreats to a shelter. Our results thus highlight that the role of female dispersal in interdemic gene flow should not be neglected.     

A female‐produced short‐range sex pheromone in the egg parasitoid Trissolcus brochymenae

The potential for short‐range sex pheromone communication by the egg parasitoid wasp Trissolcus brochymenae (Hymenoptera: Platygastridae) was investigated in closed arena bioassays. Males of this parasitoid showed more antennal drumming and more frequent mounting behaviour on 1‐ to 2‐d‐old virgin females compared with 8‐d‐old virgin females. Male copulation attempts were fewer with previously mated females than with virgin females. Males courted and made copulation attempts with 1‐ to 2‐d‐old female cadavers, but not with male cadavers or with female cadavers rinsed in organic solvents of different polarities. Male attraction to female cadavers was re‐established by treating cadavers with acetone extracts of females, but not with ether or hexane extracts. In experiments using female cadavers dissected into head, mesosoma, and gaster, and then reassembled using one unwashed body section and two body sections washed in acetone, males were attracted only to the reassembled cadavers with an unwashed mesosoma. These findings suggest that (1) courtship behaviour in males of T. brochymenae is triggered by a short‐range sex pheromone produced by females; (2) the age and the physiological condition of females (virgin/mated) influence pheromone release or production; (3) the female's mesosoma is the source of the sex pheromone; and (4) polar components of the sex pheromone play a major role in influencing male behaviour. Our results suggest that quasi‐gregarious egg parasitoids are selected for short‐range rather than long‐range sex pheromones.     

Emergence,dispersal, and mate finding via a substrate‐borne sex pheromone in the parasitoid Metaphycus luteolus

Metaphycus luteolus Timberlake (Hymenoptera: Encyrtidae) is a facultatively gregarious parasitoid of soft scale insects. We conducted behavioral experiments to better understand the mating structure of this species. Emergence of male and female offspring is synchronized, beginning at the onset of photoperiod. Both sexes are able to disperse, although dispersal of males from natal patches appears to take longer than dispersal of females. We demonstrated the presence of a female‐produced contact pheromone using open arena bioassays and motion tracking software, testing residues deposited by walking females, and extracts of females. Males responded to the females' ‘chemical footprints’ and to acetone and hexane extracts of females by searching and arrestment on the chemical residues. Responses of males were dose dependent and diminished with time since the stimulus was deposited. Our findings support the hypothesis that non‐local mating among wasps emerging from different hosts may be quite common in these parasitoids. The implications of our results for the mating structure and previously documented sex ratio patterns of these parasitoids are described.     

Experimental Validation of Sex Differences in Spatial Behavior Patterns of Free‐Ranging Snakes: Implications for Social Interactions

Social interactions often play a significant role in determining patterns of spatial use. Although snakes are generally thought of as asocial, recent spatial dispersion studies suggest that the spatial ecology of snakes may be more strongly influenced by social interactions than previously thought. We investigated the spatial behavior patterns of a western cottonmouth (Agkistrodon piscivorus) population in east Texas by uniquely combining radio‐telemetry studies on free‐ranging snakes with experimental arena trials with captive individuals from the same population. Observations from the radio‐telemetry study on free‐ranging A. piscivorus indicated that females were more gregarious than males. In the follow‐up study, spatial dispersion data from captive snakes maintained in experimental field arenas yielded similar results to spatial behavior patterns of free‐ranging individuals. When compared to random experimental controls, these data suggest that observed spatial behavior patterns are related to mechanisms associated with both active avoidance among males and conspecific attraction among females. In addition to uniquely combining both free‐ranging and captive observations, this is the first snake study to demonstrate sex differences with both of these divergent (attraction and avoidance) spatial patterns within a single population. Thus, similar to other vertebrate groups, a growing body of literature suggests that social interactions in snakes should be strongly considered in interpretations of spatial ecology and behavior.     

Characterizing phase‐related differences in behaviour of Schistocerca gregaria with spatial distribution analysis

The behaviour of locusts has been studied extensively using two approaches: (1) analysing a single individual's response to a group stimulus or (2) using group conditions to look at aggregation patterns. The second approach has, in contrast with the first one, not been improved in terms of statistical analyses since the 1960s. In the present study, we propose a spatial statistics approach of point‐pattern analysis to improve the group‐based assessment of behavioural phase characterization. This diagnostic tool was developed and tested in the laboratory with comparative analysis of solitarious (isolation‐reared) and gregarious (crowd‐reared) desert locusts, Schistocerca gregaria (Forskål) (Orthoptera: Acrididae). The spatial distribution patterns of 10 either solitarious or gregarious third‐instar hoppers were characterized with nearest neighbour distance measurements in a circular arena. The temporal sequence of spatial disposition of locusts was recorded with a digital camera taking photographs at regular intervals. The approach of point‐pattern analysis focused on the spatial distribution of observed events and allowed us to make inferences about the underlying process that generated them. The results confirmed that our diagnostic tool could identify that crowd‐reared hoppers tended to aggregate more to conspecifics than isolation‐reared ones. We could also verify that isolation‐reared hoppers were less active than crowd‐reared ones, but this was only true at the beginning of the experiments. The spatial statistics approach proposed in the present study could help with observations of phase‐related differences in the behaviour of locusts.     

Relevance of visual and olfactory cues for host location in the mustard leaf beetle Phaedon cochleariae

The relevance of visual and olfactory cues for host‐plant location is investigated in males and females of the oligophagous mustard leaf beetle Phaedon cochleariae Fabricius (Coleoptera: Chrysomelidae). Different objects are offered in a walking arena and the behaviour of beetles is observed. Beetles orient toward vertically or horizontally striped black and white pattern independent of stripe orientation. The results suggest that contrast facilitates orientation in the field, whereas the pattern itself may be less important for host location in dense vegetation. The response to green and yellow objects is tested to investigate discrimination abilities between young (green) and mature (yellow) leaves. Beetles prefer green over yellow independent of material (cardboard or leaves of Nasturtium officinale R. Br., Brassicaceae). Preference behaviour tested in a dual‐choice contact assay coincides with visual preferences, where adults prefer young, more nutritious leaves for feeding and oviposition. Furthermore, females discriminate between visual cues of green leaves and green cardboard, whereas males do not, indicating that females are more sensitive in colour discrimination. Differences in colour wavelength influence the choice of beetle behaviour more strongly than differences in intensity. Both sexes of P. cochleariae prefer volatiles of the host plant N. officinale, whereas only females respond to the main volatile compound 2‐phenylethyl isothiocyanate. Given a choice between visual and olfactory cues, males orientate towards the colour cues, whereas females do not show any preferences. In males, visual cues may thus override olfactory cues, whereas, in females, both are equally important, which may reflect different ecological requirements and/or physiological abilities.     

Widespread 'hilltopping' in Acraea butterflies and the origin of sex-role-reversed swarming in Acraea encedon and A. encedana

In some populations of the butterflies Acraea encedon and A. encedana, most females are infected with a bacterium that kills their sons. The resulting shortage of males is associated with females adopting a sex‐role‐reversed mating system, in which females swarm at landmarks such as hilltops and compete for males. We have observed the mating behaviour of Acraea species that are not known to be infected with the male‐killer. In over half of these species, males were found to aggregate on hilltops. It is likely that this behaviour was ancestral to the sex‐role‐reversed swarms of Acraea encedon and A. encedana, and we discuss how the spread of the male‐killing infection may have converted this mating system into sex‐role‐reversed swarming.     

Small-scale vegetation patterns in the parental environment influence the phase state of hatchlings of the desert locust

Desert locusts (Schistocerca gregaria Forskål (Orthoptera: Acrididae)) change phase in response to population density. Solitarious insects avoid one another; when crowded, they shift to the gregarious phase and aggregate. Laboratory experiments and individual‐based modelling have shown that small‐scale resource distribution can affect locust phase state via an influence on crowding. Laboratory work has also shown that parental phase state is transmitted to offspring via maternal inheritance. These effects had not been investigated in the field previously. We maintained small populations of adult desert locusts in semi‐field enclosures with different distribution patterns of a single plant species (Hyoscyamus muticus L. (Solanaceae)). The offspring of locusts exposed to more clumped patterns of vegetation exhibited more gregarious behaviour when tested in a behavioural phase assay than did progeny from parents left in enclosures with more scattered vegetation. These effects on nymphal behaviour appeared to be mediated by influences of resource distribution on adult phase state. Phase state in small semi‐field populations was influenced by small‐scale vegetation distribution. Phase differences engendered by environmental structure were maintained in time and transmitted to progeny.     

The effect of nest aggregation on the reproductive behaviour of the peacock blenny Salaria pavo

The effect of nest aggregation in courtship behaviour was tested experimentally in an ecologically constrained, sex-role reversed population of the peacock blenny Salaria pavo . Mixed sex groups of eight males and eight females were tested in experimental tanks, containing eight potential nests either aggregated or dispersed. In the aggregated treatment, males spent more time inside their nests and monopolized other potential nests, causing a female-biased operational sex ratio (OSR). In the aggregated treatment, females also expressed more courtship behaviour. The results in general support the prediction that the aggregation of nests promotes male monopolization of potential nests, resulting in fewer nest-holding males and therefore a female-biased OSR that leads to the reversal of sex roles.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

Female Attraction to Conspecific Chemical Cues in the Palmate Newt Triturus helveticus

Although chemosignals are largely used in sexual communication in urodeles, olfactometer studies in newts provided contrasting results about the sex specificity of female behavioural responses. Because long‐range sexual advertisement is believed to be costly, some species might restrain this activity to close interactions with conspecifics. We tested chemical‐mediated sexual attraction in female palmate newt (Triturus helveticus) by measuring the attraction to male and female odours in a linear water olfactometer. Unexpectedly, females were attracted towards conspecifics regardless of sex. They did not show attraction towards Limnaea stagnalis, a common sympatric aquatic gastropod. These results do not support the use of long‐range male sexual signalling in the palmate newt. Instead, conspecific attraction is likely to promote aggregation of males and females in breeding ponds. Observations in the field and in the laboratory tend to support the aggregative behaviour of this species. We discuss the possible function of conspecific attraction in this context. Heading towards any conspecific would increase the probability of finding potential mates. Chemical cues do not need to be sex‐specific at that stage so that long‐range sexual advertisement might be unnecessary. This work emphasizes the need for studies investigating the evolutionary relationships between sexual signalling systems and population‐distribution patterns.     

The role of functional constraints in nonrandom mating patterns for a dance fly with female ornaments

Most hypotheses to explain nonrandom mating patterns invoke mate choice, particularly in species that display elaborate ornaments. However, conflicting selection pressures on traits can result in functional constraints that can also cause nonrandom mating patterns. We tested for functional load‐lifting constraints during aerial copulation in Rhamphomyia longicauda, a species of dance fly that displays multiple extravagant female‐specific ornaments that are unusual among sexual traits because they are under stabilizing selection. R. longicauda males provide females with a nuptial gift before engaging in aerial mating, and the male bears the entire weight of the female and nuptial gift for the duration of copulation. In theory, a male's ability to carry females and nuptial gifts could constrain pairing opportunities for the heaviest females, as reported for nonornamented dance flies. In concert with directional preferences for large females with mature eggs, such a load‐lifting constraint could produce the stabilizing selection on female size previously observed in this species. We therefore tested whether wild‐caught male R. longicauda collected during copulation were experiencing load‐lift limitations by comparing the mass carried by males during copulation with the male's wing loading traits. We also performed permutation tests to determine whether the loads carried by males during copulation were lighter than expected. We found that heavier males are more often found mating with heavier females suggesting that whereas R. longicauda males do not experience a load‐lift constraint, there is a strong relationship of assortative mating by mass. We suggest that active male mate choice for intermediately adorned females is more likely to be causing the nonrandom mating patterns observed in R. longicauda.     

The mating system of the true fruit fly Bactrocera tryoni and its sister species,Bactrocera neohumeralis

The frugivorous “true” fruit fly, Bactrocera tryoni (Queensland fruit fly), is presumed to have a nonresourced‐based lek mating system. This is largely untested, and contrary data exists to suggest Bactrocera tryoni may have a resource‐based mating system focused on fruiting host plants. We tested the mating system of Bactrocera tryoni, and its close sibling Bactrocera neohumeralis, in large field cages using laboratory reared flies. We used observational experiments that allowed us to determine if: (i) mating pairs were aggregated or nonaggregated; (ii) mating system was resource or nonresource based; (iii) flies utilized possible landmarks (tall trees over short) as mate‐rendezvous sites; and (iv) males called females from male‐dominated leks. We recorded nearly 250 Bactrocera tryoni mating pairs across all experiments, revealing that: (i) mating pairs were aggregated; (ii) mating nearly always occurred in tall trees over short; (iii) mating was nonresource based; and (iv) that males and females arrived at the mate‐rendezvous site together with no evidence that males preceded females. Bactrocera neohumeralis copulations were much more infrequent (only 30 mating pairs in total), but for those pairs there was a similar preference for tall trees and no evidence of a resource‐based mating system. Some aspects of Bactrocera tryoni mating behavior align with theoretical expectations of a lekking system, but others do not. Until evidence for unequivocal female choice can be provided (as predicted under a true lek), the mating system of Bactrocera tryoni is best described as a nonresource based, aggregation system for which we also have evidence that land‐marking may be involved.     

Ladies last: diel rhythmicity of adult emergence in a parasitoid with complementary sex determination

Protandry, the earlier adult emergence of males, is explained as either an adaptive strategy maximizing male mating opportunities at the same time as minimizing female pre‐reproductive mortality, or as an incidental by‐product of sexual dimorphism fuelled by selection for other life‐history traits. Adult emergence sequences are monitored of broods of the gregarious larval endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) undergoing pupal development under different temperature regimes. As a haplodiploid species with single‐locus complementary sex determination, gender in C. glomerata is determined by the genotype at one sex locus. Haploids are always male, whereas diploids are female when heterozygous but male when homozygous at the sex locus. Sibling mating promotes homozygosity and thus the production of diploid males. Diploid males are produced at the expense of females, and impose a genetic burden on individuals and populations, despite their exceptional fertility in C. glomerata. Emergence of broods is typically completed within 2 days. Irrespective of temperature, males emerge earlier and within a shorter time interval than females, and a majority of the males in a cluster emerge before the first female. The implications of an incomplete temporal segregation of the sexes on the incidence of inbreeding in C. glomerata are discussed in the light of its sex determination mechanism and its patterns of mating, host exploitation and natal dispersal.     

Sexual attractiveness shared by both sexes mediates same‐sex sexual behaviour in the parasitoid wasp Telenomus triptus

The mating behaviour of a quasi‐gregarious egg parasitoid Telenomus triptus Nixon (Hymenoptera: Platygastridae), which exploits egg masses of a stink bug Piezodorus hybneri (Heteroptera: Pentatomidae), is examined in the laboratory. In this parasitoid wasp, male adults that emerge earlier stay at the natal egg mass and mate with subsequently emerging females. In the present study, a male adult that encounters the emergence of another male always waits for it to egress, and then mounts the newly emerging male. To examine why males of T. triptus show same‐sex sexual behaviour, male adults are presented with a parasitized host egg mass or a freshly killed wasp. Male adults are observed to remain at host egg masses from which only male wasp(s) had emerged. In addition, male adults attempt to copulate with freshly killed young male wasps. It is suggested that newly emerging male wasps are targets of same‐sex sexual behaviour because they possess cues for male sexual behaviour similar to the cues of females. Both the sex and age of freshly killed wasps affect the frequency of the sexual behaviour of male adults: females are more attractive than males, although their attractiveness declines with age. When the mating opportunity is restricted to the natal egg mass, the costs of failing to notice newly emerging female adults should be extremely high. Therefore, males are forced not to discriminate the sex, resulting in same‐sex sexual behaviour.     

The ecology of sexual selection: Mean crowding of females and resource-defence polygyny

Summary Resource-defence polygyny is a mating system that offers an opportunity for strong sexual selection. I express the degree of aggregation or clustering of females at resources at the time of mating using Lloyd's measures of microspatial aggregation, mean crowding (m ^*) and patchiness (P). Here, I show that Lloyd's two ecological measures of aggregation can be used to quantify the relationship between resource-based polygyny and sexual selection. The advantage of using these measures is that some of the statistical machinery for estimating the degree of mean crowding and testing for density dependence in comparative ecological studies can be applied to estimating the strength of sexual selection and comparing different mating systems. When females vary in their tendency to aggregate with other females and males vary from one another in their ability to guard aggregations of females, a run-away process of sexual selection is possible. The largest aggregations of females have a higher proportion of the females most prone to aggregate or most tolerant of crowding. The males with the highest mating success have daughters more prone to aggregate and sons better at defending aggregated females against other males. As a result, females are selected to over-aggregate with respect to resources. The run-away process halts when single males cannot defend and maintain exclusive reproductive access to clusters of females as evidenced by extraharem copulations and paternity.     

How Do Females’ Genetic Characteristics Influence Male Mate Preference in the Terrestrial Isopod Armadillidium vulgare?

Genetic diversity is a key factor that can influence mate choice in many species. We experimentally determined the influence of this factor on mate preference in the crustacean terrestrial isopod Armadillidium vulgare. This biological model is gregarious which could increase the risk of inbreeding by mating with closely related partners. Mechanisms of inbreeding avoidance during mate choice can thus be expected. Moreover, previous studies predict that males would be the choosy sex. We performed Y‐choice tests giving males the choice between two females presenting different levels of heterozygosity and genetic similarity to the male. Our results show potential inbreeding avoidance according to the genetic characteristics of females presented to males. The higher the variation in genetic similarity to the male between females is, the higher the preference of the male towards the most dissimilar female is. Hence, male preferences may only be detectable when the difference between females’ genetic characteristics is large enough. If heterozygosity is associated with fitness in A. vulgare (as in many organisms), the patterns of mate preference we observe may be adaptive.     

Evidence for male- and juvenile-specific contact pheromones of the common bed bug Cimex lectularius

Males and females of the common bed bug, Cimex lectularius L. (Heteroptera: Cimicidae), have been shown to produce and respond to an aggregation pheromone. We tested whether juvenile C. lectularius also produce and respond to aggregation pheromone, and whether the pheromone is perceived by contact chemoreception. In dual‐choice laboratory experiments, juveniles, but not males or females, preferred juvenile‐exposed paper discs to control discs. Unlike juveniles, males and females preferred male‐exposed paper discs to control discs. Neither juveniles, males, nor females preferred female‐exposed discs to control discs. When test stimuli were inaccessible, C. lectularius failed to show any preference. Male‐ and juvenile‐specific contact pheromones may have contrasting functions of marking shelters as safe refugia for development and growth (juveniles) or mate encounter (adults), but result in the same phenomenon, the aggregation of conspecifics.