Feeding ecology and evidence for amino acid synthesis in the periodical cicada (Magicicada)

The periodical cicadas of the genus Magicicada (including M. septendecim, M. cassini, and M. septendecula) have the longest juvenile life span of any insect, living underground for 13 or 17 years and feeding exclusively on root xylem fluids. Due to their inaccessible life cycles very little is known about cicada nutrition, despite the fact that members of Magicicada can achieve a very large biomass in woodland habitats east of the Mississippi and hence constitute a major part of the ecosystem where they occur in high densities. Live cicadas were collected at two sites in early June of 2004, during the emergence of Brood X (both M. septendecim and M. cassini were recovered). We used a combination of stable isotopic measurements (δ¹⁵N and δ¹³C) and multivariate statistical techniques to test for differences in resource acquisition among the cicada species and sexes collected at two locations within the 17-year periodical Brood X range. The amino acid constituents of cicada chitin and organs, plus xylem extracted from a deciduous sapling, were also analyzed. The data show that male and female cicadas have different carbon fractionations, which could reflect differential resource utilization due to oviposition in females. Several essential amino acids for the cicada were absent in xylem. Carbon-isotopic composition of all amino acids in the cicadas was distinctly different from the limited set measured in the xylem. Because of the differences in isotopic composition, we conclude that amino acids were synthesized de novo rather than incorporated directly, most likely produced by endosymbiotic bacteria.     

The role of auditory stimuli in the courtship of Drosophila melanogaster

Simulated courtship song of male Drosophila melanogaster was played to males or females of this species. Upon receiving the song males increase their locomotor activity and start courting each other, whereas females reduce their locomotor activity. In wingless males the locomotor activity difference between the silent control and the experimental sound situation is much larger than in winged males, due to the inactivity of wingless males in the control situation. Males which had been kept singly up to the time of the experiment exhibit higher locomotor and sexual activity than group housed males. A second component of the male courtship song ‘sine song’ is described, together with experiments which investigate the sensory basis of the effect male courtship song has on males.     

Mate recognition in members of the Drosophila nasuta complex

An examination of the courtship patterns of species belonging to the Drosophila nasuta complex shows that markings which are characteristic of males are almost certainly involved in their recognition as conspecifics by females. In each species the males perform a courtship sequence in such a way as to expose these markings to the female. A phylogeny of the species based on both male courtship patterns and the associated male markings is constructed. This phylogeny is compared to previously published phylogenies of the same group which were based on fixed and floating chromosomal differences between the species. Finally, the evidence which the complex provides for the utility and general significance of a new species concept, the Recognition Concept, is evaluated.     

Female and male interactions during courtship in Calopteryx maculata and C. dimidiata (Odonata: Calopterygidae): Influence of oviposition behaviour

Calopteryx maculata and C. dimidiata damselfly females respond to male courtship with specific displays which signal differences in their receptivity. These include a rejection (wing spreading) and an invitation (wing-flipping) display, as well as a neutral (sit still) response. There are interspecific differences in the likelihood of each female display and in male responses to these displays. C. maculata males persist in courtship irrespective of female response, while C. dimidiata males generally stop courting when the female's response is rejection or neutrality. I suggest that these differences result from interspecific differences in oviposition behaviour. Female C. maculata oviposit at the water surface, which exposes them to disturbance by males attempting to mate. Females are therefore likely to remate to secure postcopulatory guarding when changing oviposition sites and males are expected to be persistent in courtship. Female C. dimidiata submerge to oviposit, which frees them from male disturbance and means that males have less control over female access to oviposition sites. Males therefore have less influence on mating by females and are expected not to persist in courtship of non-receptive females.     

High-frequency clicks of Hawaiian picture-winged Drosophila species

In this study, it is shown that the males of several picture-winged Drosophila subgroup species produced high-frequency clicking sounds when courting females. At the beginning of the courtship, the males may semaphore or vibrate their wings with a large amplitude, producing no audible sounds. After these ‘preliminary’ wing vibrations the males set their wings backwards in a normal resting position and vibrate them with a small amplitude, producing loud clicking sounds (up to 15000 cps), which differ from all Drosophila sounds described so far in both their spectral and their temporal structure. When producing these sounds the males always touch the abdomen of the female with their front legs, which might help the females receive the sounds as vibrational signals.     

Comparative courtship behaviour in two species of the parasitic chalcid wasp Melittobia (Hymenoptera: Eulophidae)

Timing and form of courtship behaviour elements constitute a major isolating mechanism for two morphologically and ecologically similar parasitic wasps. Because males are blind, the complex courtship of Melittobia chalybii and a new as yet undescribed species previously confused with it, designated as M. sp. A, depends upon tactile and chemical cues. In both species, after the female orients to the male's abdomen, the male turns in the direction of contact and mounts. However, chalybii males antennate females continuously, lifting mesothoracic legs at regular intervals, while in sp. A male antennation alternates with metathoracic leg pumping. Courtship and copulation duration are greater in Chalybii than in sp. A.     

Sexual behaviour of a female sterile mutant of Drosophila melanogaster

The courtship of male Drosophila melanogaster to genetically sterilised females (homozygous for the fs(2)B gene) and their fertile siblings (heterozygous for fs(2)B) was investigated. Courtship measures were made with females from two dietary conditions (‘normal’ and ‘glucose’) and of two reproductive states (virgin; inseminated). Females homozygous for the mutant gene and females maintained on glucose were found to have a lower level of sexual receptivity. The results also show that these females received more intensive courtship than the control flies. The effects of insemination differed in their extent dependent upon genotype and diet. Locomotor activity was markedly affected by diet and to a lesser extent by genotype. The differences observed in courtship behaviour cannot be wholly attributed to differences in activity level. The similarity of the effects of the glucose diet and female sterility suggest that the ovaries, or neuroendocrine factors linked in feedback relationships to them, are responsible for the observed effects.     

Courtship-stimulating volatile compounds from normal and mutant Drosophila

Volatile compounds from Drosophila melanogaster males and females dramatically affect male courtship behaviour. These substances, which have been extracted from flies of different ages and genotypes, have been analysed by gas chromatography (GC) and in behavioural assays. Extracts from virgin females and males have different gas chromatographic profiles, which may reflect the fact that extract from virgin females stimulates high levels of courtship between males over short distances, while extract from mature wild-type males does not affect sexual behaviour. However, volatile compounds from very young males or males expressing the fruitless (fru) mutation do stimulate courtship between males, and chromatographic profiles of young male and fru male extracts differ from the GC profile of extracts from mature wild-type males.     

Mating behaviour in the Australian reptile tick Aponomma hydrosauri

Females of the Australian reptile tick Aponomma hydrosauri make few movements once they are attached to a host. Males move significantly more often on hosts with females attached. Male movement often leads to contact and courtship with females. Movement is most frequent in the early part of the dark phase of a photoperiod. After contact the courtship, both on the host and in experimentally induced off-host situations, follows a stereotyped sequence of positional orientations, here divided into six phases. Copulation does not automatically follow courtship initiation, and in particular successful courtship requires females to respond to male initiation by lifting their hind legs to allow the male to crawl under. Compared to other animal species, the males of A. hydrosauri play an unusually passive role in mate seeking. This may be explained by the extremely patchy distribution of the mate-finding environment, lizards.     

Differential responses of yellow-breasted chats, Icteria virens, to male and female conspecific model presentations

A detailed understanding of the behavioural interactions between males and females is crucial for elucidating the selection pressures shaping mating system evolution; however, these interactions are often difficult to observe, particularly in free-living populations. We simulated extraterritorial intrusions by presenting conspecific models on to territories of a Neotropical migratory passerine, the yellow-breasted chat, Icteria virens. Simulated intrusions elicited different responses based on the sex of the focal individual and the sex and species of the model. Behavioural responses of the focal individuals were largely directed towards conspecific models, whereas simulated heterospecific intrusions, using a Carolina wren, Thryothorus ludovicianus, model, elicited minimal response. Male song was not related to any particular model presentation. Males directed mating behaviours, including courtship displays and copulations, exclusively towards the female conspecific model. Males behaved aggressively towards the conspecific male model. In contrast, females never showed any courtship behaviour towards the male model and showed significant aggression towards the conspecific female model. The results of this study reveal that territorial aggression in yellow-breasted chats is strongly intrasexual. Additionally, simulated female extraterritorial intrusions, but not male intrusions, resulted in extrapair courtship and copulation. This pattern is likely to be typical of many socially monogamous species and points to the behavioural mechanisms underlying both monogamy and extrapair mating systems.     

A meal or a male: the ‘whispers’ of black widow males do not trigger a predatory response in females

Introduction

Female spiders are fine-tuned to detect and quickly respond to prey vibrations, presenting a challenge to courting males who must attract a female’s attention but not be mistaken for prey. This is likely particularly important at the onset of courtship when a male enters a female’s web. In web-dwelling spiders, little is known about how males solve this conundrum, or about their courtship signals. Here we used laser Doppler vibrometry to study the vibrations produced by males and prey (house flies and crickets) on tangle webs of the western black widow Latrodectus hesperus and on sheet webs of the hobo spider Tegenaria agrestis. We recorded the vibrations at the location typically occupied by a hunting female spider. We compared the vibrations produced by males and prey in terms of their waveform, dominant frequency, frequency bandwidth, amplitude and duration. We also played back recorded male and prey vibrations through the webs of female L. hesperus to determine the vibratory parameters that trigger a predatory response in females.

Results

We found overlap in waveform between male and prey vibrations in both L. hesperus and T. agrestis. In both species, male vibrations were continuous, of long duration (on average 6.35 s for T. agrestis and 9.31 s for L. hesperus), and lacked complex temporal patterning such as repeated motifs or syllables. Prey vibrations were shorter (1.38 - 2.59 s), sporadic and often percussive. Based on the parameters measured, courtship signals of male L. hesperus differed more markedly from prey cues than did those of T. agrestis. Courtship vibrations of L. hesperus males differed from prey vibrations in terms of dominant frequency, amplitude and duration. Vibrations of T. agrestis males differed from prey in terms of duration only. During a playback experiment, L. hesperus females did not respond aggressively to low-amplitude vibrations irrespective of whether the playback recording was from a prey or a male.

Conclusions

Unlike courtship signals of other spider species, the courtship signals of L. hesperus and T. agrestis males do not have complex temporal patterning. The low-amplitude ‘whispers’ of L. hesperus males at the onset of courtship are less likely to trigger a predatory response in females than the high-amplitude vibrations of struggling prey.     

Mating systems and sex ratios in the egg parasitoids, Trichogramma dendrolimi and T. papilionis (Hymenoptera: Trichogrammatidae)

Arrhenotokous parthenogenesis was confirmed in Trichogramma dendrolimi and T. papilionis. Subsequently the possible links between mating systems and biological traits of males, and sex ratios, were investigated in these two species, using Papilio xuthus eggs for their hosts. T. dendrolimi males attained 100% insemination of females in the parasitized host before egress from it. The high insemination rate was guaranteed by male precedence in emergence, lack of courtship in mating behaviour and short copulation time, combined with a long stay of emerging wasps within the host. The males were often the last to leave a host and made no mating attempts outside the host. Most but not all T. papilionis females were also inseminated in the host. The lower insemination rate of this species resulted from almost simultaneous emergence of both sexes, which prevented males from mating with inactive females. Another mating site of T. papilionis was just outside the host, which males left prior to most females. A reduced possibility of outbreeding was inferred in T. dendrolimi on the grounds that males were short-lived, frequently failed to expand their wings and died in the host. The reduced outbreeding was reflected in considerably lower sex ratios in T. dendrolimi.     

Delayed courtship in the fiddler crab Uca musica terpsichores

Courting male fiddler crabs of the species Uca musica terpsichores congregate in the upper central portion of the colony, while receptive females leave their burrows located at the colony's periphery and wander among the communally displaying males prior to choosing a mate. I observed that courting males in a newly-established population were significantly smaller than courting males in large high-density colonies. This observation led to a series of translocation experiments designed to ascertain whether high population density influences the size (=age) at which males begin to court. Smaller courting males from a low-density population failed to court after being placed among larger courting males in a high-density population. The reciprocal translocation revealed that smaller noncourting males from the high-density population would start courting shortly after being placed in a low-density population. Smaller males placed in the high-density population were subsequently observed significantly further away from where they were initially placed than were larger males similarly translocated. The results suggest that smaller males delay courtship activities once they are forced, via encounters with larger males, to the periphery of the colony. I believe that both intrasexual selection (competition from larger males) and intersexual selection (female choice of large males) are responsible for the delay in male courtship activities.     

Male displays and female preferences in the courtship of a gregarious cricket

The courtship of males of the gregarious cricket Amphiacusta maya involves a variety of signals. The quantitative aspects of both successful and unsuccessful courtship sequences were examined to determine whether certain aspects of male displays were correlated with female mating preferences regardless of which male performed them. Although variability among males was high for most courtship components measured, I found no evidence of female choice with respect to the courtship variables studied. About 35% of the duration of each male courtship sequence is devoted to chirping, but there were no differences in either the likelihood of copulation or the latency to copulation between normal males and experimentally silenced males. The possibility that intrinsic differences in male quality explained the variability in courtship duration was examined with a two-way analysis of variance. The variance in courtship duration was attributable to variance among females, not to variance among males. Thus the courtship behaviour of male A. maya is variable enough to allow females to exert stabilizing or disruptive selection on displays, but there is no evidence that females use the available information.     

Short‐Term Nutritional Limitation Affects Mating Behaviour and Reproductive Output in Dwarf Spiders

Mating decisions can vary considerably depending on individual experience, mate availability and nutritional status. Here, we applied short‐term dietary restrictions to adult female spiders that were well fed during the juvenile stage in an effort to understand whether and how brief periods of food shortage can influence male and female mating decisions and mating behaviour. To assess whether responses vary between closely related species, we conducted the same experiment on the dwarf spiders Oedothorax retusus and O. apicatus. During courtship and mating, males of both species offer secretion to females from glandular tissue in their prosoma. Females were subject to food shortage over a period of 3 wks (‘low‐diet’ treatment, LD) or fed regularly (‘high‐diet’ treatment, HD). We compared courtship probability, mating probability/behaviour, and reproductive output between dietary groups and species. In both Oedothorax species, females in the LD treatment were less likely to mate and more aggressive towards males. Furthermore, LD females produced egg sacs that were significantly lighter than were those of the HD females. Effects of food deprivation on copulation duration, gustatory behaviour and oviposition latency differed between species. Our study shows that short periods of dietary restriction during the adult stage can strongly affect mating behaviour and reproductive output with differences between closely related species.     

Decapitated females as a tool in the analysis of Drosophila behaviour

Decapitated females provide a method of arranging variable stimulus environments to evaluate components of behaviour involved in sexual and frequency dependent selection. Conspecific matings are bypassed via male preference for normal females per se. Presence of conspecific decapitatated females induces male to increase insemination of alien females. Discrimination mechanisms, including sign stimuli, are the least labile, and excitability is the most labile component of courtship. D. subpalustris males with abnormal stimulation use a qualitatively different courtship element not present in their normal repertoire. An efficiency index (courtship/insemination) shows D. palustris males, with more heterogenous stimulation in their courtship, more effective in overcoming female reluctance. Sexual isolation may differ operationally from sexual selection in the necessity for particular stimuli rather than more stimulation.     

An observational study of scent-marking in coyotes, Canis latrans

Urination and defaecation patterns of free-ranging coyotes (Canis latrans) were studied in the Grand Teton National Park, Jackson, Wyoming, for two years. The vast majority of urinations by adult males and females were involved in ‘marking’, and differentiating between ‘marking’ and ‘elimination’ may not be necessary. Our results may be summarized as follows: (1) raised-leg urinations (RLU) performed by males were most frequently used in marking. (2) Females marked throughout the year using the squat (SQU) posture. (3) Snow tracking and reading snow sign resulted in a gross underestimate of the relative frequency of SQU's and a large overestimate in the relative frequency of defaecations (DEF) when compared to results obtained by direct observation. (4) There was sexual dimorphism for the contexts in which marking occurred. Overall, marking by males was associated with courtship and mating, with travelling, and with aggression. Marking by females was associated with the acquisition and possession of food and with the denning season. (5) Marking rates per coyote increased in groups larger than two animals. (6) RLU marking rates were greatest in areas of high intrusion when compared to denning areas and areas in which non-group members infrequently tresscent odours are important in orienting individuals in space but do not represent in and of themselves barriers to movement.     

Evolutionary significance of courtship conditioning in Drosophila melanogaster

Males of the fly Drosophila melanogaster initially court mated, unreceptive females but later develop an avoidance reaction against them and even become temporally unresponsive to virgin females. This conditioned inhibition has been described as an associative process, the conditioned stimulus being a mixture of pheromones on the female's cuticle. To assess the evolutionary significance of courtship conditioning we recorded and analysed the male's behaviour during the conditioning process. The time traces of individual males were marked by an abrupt behavioural change. The time he spent courting suddenly decreased, and the frequency of ‘turn-away’ events at the same time sharply increased. Thus, the gradual decline of courtship observed as a group average does not reflect a slow change in motivation of the individual male but rather the interindividual differences of an active, experience-guided all-or-none decision to stop courting and to avoid the female. Three recently collected D. melanogaster strains were each maintained under two distinct mating conditions. Males were kept together with females for either ca. 2 weeks or 18 h. After 21 generations males of the two regimes differed markedly in their behaviour towards mated females. With long interaction periods between males and females, selection favoured courtship conditioning, while with short periods no such selection was observed. Slowly recovering receptivity of mated females may be needed for the maintenance of courtship conditioning. Courtship conditioning in D. melanogaster appears to be a fitness-relevant behaviour adapted to high-density populations with females mating a second time.     

FUNCTIONAL ASPECTS OF DROSOPHILA COURTSHIP

1. The elements that make up the courtship behaviour of males and of females are briefly described. It is pointed out that some of the terms used, such as female ‘repelling’ behaviour, are misleading as they do not reflect the known functions of the behaviours. 2. Evidence has been presented for a number of distinct pheromones with different functions during courtship. These claims are critically examined as the evidence is incomplete and at times conflicting. It seems unlikely that any pheromones other than those acting over a very short distance are involved in courtship. There is sound evidence for an aphrodisiac pheromone produced by all females which stimulates male courtship. A pheromone, which may be the same one, is produced by males less than 12 h old, which also stimulates male courtship. No function is ascribed to this pheromone. Fertilized females either produce less aphrodisiac pheromone or they may, in addition, produce one that inhibits male courtship. Mature males may also produce an inhibitory pheromone. Females produce a contact pheromone which is species-specific and involved in sexual isolation. It is not at present clear whether this is different from the aphrodisiac pheromone. 3. There is considerable variability in the importance of vision in courtship. Many species will mate satisfactorily in the dark, suggesting that visual stimuli are not critical. Most species use vision to orient towards one another and for males to track and follow females. Even in light-independent species such as D. melanogaster, specific visual signals may be used in courtship although they are not obligatory. Thus the red eye of the male is a sexual signal for females. Conversely, some light-dependent species do not appear to make use of visual signals as a major factor in courtship. Some, however, do perform behaviours that are clearly visual and which may act to emphasize markings on wings, head or body. 4. The majority of Drosophila species perform courtship songs by vibrating one or both wings. The songs produced by males sexually stimulate the females. They are species specific and there is considerable indirect and some direct evidence that the songs are involved in sexual isolation. Males of many species produce two different songs during courtship and it is probable that one is concerned mainly with sexual stimulation and the other with species recognition. Females of certain species of Drosophila and Zaprionus also sing during courtship and these songs may aid species recognition by males. In addition males and unreceptive females perform ‘aggressive’ songs. 5. Almost all studies of Drosophila courtship have been made in very confined conditions in the laboratory. Interpretation of some of the results obtained in this way may require modification in the light of ecological research and observation of courtships under more natural conditions.     

The chemical basis for sex recognition in Drosophila melanogaster

Chemical cues were recognized to play a predominant role in initiating male courtship behaviour in Drosophila melanogaster as measured by the number and duration of wing-vibration responses elicited in test males. The effect was associated with compounds specific to the female cuticle, for which we describe a simple extraction procedure. Female active extracts were compared with behaviourally non-active extracts from males, using gas-liquid and thin-layer chromatography. Using these preparative methods, long-chain hydrocarbons were isolated and activity was found only among unsaturated molecules. One, heptacosadiene, inducing the highest level of courtship, appears to be the main aphrodisiac pheromone of the female D. melanogaster. This compound is specific to females of the species and is the most abundant of their cuticular hydrocarbons.