昆虫求偶行为在分类学上的应用

昆虫求偶行为是两性间相互吸引并完成各种生殖活动的一种定向行为,其形式多样,存在种的特异性。不同物种其求偶行为不同,以此来帮助区分特定类群近缘种、识别生物学种显得非常有意义。本文综述了性信息素求偶行为、鸣声求偶行为、闪光求偶行为以及献礼求偶行为在分类学上的应用。     

Male competition is disruptive to courtship in the Pacific blue‐eye

The present study examined the effects of competition on male courtship in the Pacific blue‐eye Pseudomugil signifer , a species of fish where females have previously been shown to use courtship, but not male fighting prowess, as an important mate choice cue. Courtship bouts directed to a stimulus female were shortest when two males were allowed to freely interact (contact treatment) and longest when there was only one male (non‐interaction). Courtship length in trials where one of two males was confined to a clear cylinder (visual) was intermediate between the other two treatments. Courtship in visual and contact treatments was constantly disrupted. The percentage of interrupted courtships was higher for the contact compared to visual treatment where aggressive interactions were also longer in duration and took place more frequently. Within contact trials, dominant males courted longer than subordinates despite both males experiencing comparable rates of courtship disruption. These results suggest that male‐male competition can have important implications for adaptive female choice particularly in circumstances where the benefits being sought by females are unrelated to male fighting ability.     

The courtship behaviour of the pipefish Nerophis lumbriciformis: reflections of an adaptation to intertidal life

The courtship behaviour of Nerophis lumbriciformis (Pisces: Syngnathidae) consists of three distinct phases (initial courtship, spawning, and embrace) marked by prominent behavioural changes. The first courtship phase is characterised by female quivering. Courtship activity increases from low to high levels of quivering, causing undulatory head movements in the female. In the second phase, the female transfers her eggs onto the male incubating ventral surface. During the final phase, the male wraps his body around the female (embrace). Females actively initiate courtship indicating that these pipefishes are courtship role reversed. The complete lack of swimming and vertical movements in the courtship behaviour of N. lumbriciformis, unlike in the behaviour of other syngnathid species, suggests an adaptation to intertidal conditions.     

Evolution and function of multimodal courtship displays

Courtship displays are behaviours aimed to facilitate attraction and mating with the opposite sex and are very common across the animal kingdom. Most courtship displays are multimodal, meaning that they are composed of concomitant signals occurring in different sensory modalities. Although courtship often strongly influences reproductive success, the question of why and how males use multimodal courtship to increase their fitness has not yet received much attention. Very little is known about the role of different components of male courtship and their relative importance for females. Indeed, most of the work on courtship displays have focused on effects on female choice, often neglecting other possible roles. Additionally, a number of scientists have recently stressed the importance of considering the complexity of a display and the interactions between its different components in order to grasp all the information contained in those multimodal signals. Unfortunately, these methods have not yet been extensively adapted in courtship studies. The aim of this study was to review what is currently known about the functional significance of courtship displays, particularly about the role of multimodality in the courtship communication context. Emphasis is placed on those cases where a complete picture of the communication system can only be assessed by taking complexity and interaction between different modalities into account.     

Changes in courtship behaviour following rejection: The influence of female phenotype in Drosophila melanogaster

Courtship can be costly and so selection should favour individual males that reduce courtship towards female types that have a low probability of resulting in copulation. One way males can do this is by associating previous courtship failure with the traits of particular rejecting females. We characterised changes in male Drosophila melanogaster courtship behaviour following a failed mating attempt with one of the four female phenotypes that varied in size, age or mating status. To do this, we assessed individual courtship behaviour for each male presented again with a female of the same phenotype that previously rejected him. Males reduced subsequent courtship most strongly for recently mated (sexually non‐receptive) females. More interestingly, males also significantly reduced courtship activity following a failed mating experience from old females but did not do so for control (large, young, virgin) or small females. As such, males significantly reduced courtship towards both female types possessing chemical cues associated with their phenotype (age and mating status), but not towards a female phenotype based on physical characteristics (body size). Our results suggest that males are able to modify their courtship behaviour following experience, but that they are better prepared to associate chemical traits that may be more reliable indicators of the likelihood of courtship failure.     

Species and sex-specific songs and courtship behaviour in the genus Zaprionus (Diptera-Drosophilidae)

Courtship behaviour and songs of six Zaprionus species show differences not only between species but also between sexes. Courtship behaviour differs from that in the related genus Drosophila. In most species, males produce two song types which may alternate or be repeated. Singing occurs during courtship displays, mounting, after copulation and briefly during male-to-male interactions. Females produce a loud whine and body rocking movements in refusal but have a species-specific pulsed song produced during normal courtship which differs from the conspecific male song. One species produces an irregular male song but a regular female song. Sex-specific songs may have selective advantages but pose problems for the sensory template hypothesis.     

Comparison of black-footed ferret (Mustela nigripes) and domestic ferret (M. putorius furo) courtship activity

Because of the scarcity of the endangered black-footed ferret (Mustela nigripes) and the amount of knowledge necessary for their conservation, surrogate research can play an important role in recovery. In this paper, we investigate surrogate behavioral research potential by comparing courtship behavior of the black-footed ferret to the congeneric domestic ferret (M. putorius furo). Ten female domestic ferrets were bred to five male domestic ferrets and eight female black-footed ferrets were bred to five black-footed ferret males. Courtship activities were defined, analyzed, and quantitatively compared between both groups. Lag sequential analysis of was used to prepare the behavioral matrices, and matrix cells were compared between groups with an equality of proportions test. Courtship patterns did not differ significantly between the two closely related species, and the domestic ferret would probably be an adequate surrogate for reproductive behavior research on the black-footed ferret.     

Comparative study of courtship in twelve phycitine moths (Lepidoptera: Pyralidae)

The courtship behavior of 12 phycitine moths (Lepidoptera: Pyralidae) was studied using frame-by-frame analysis of video recordings. Behavioral transitions during courtship were quantified for selected species and kinematic diagrams of courtship sequences were constructed. Interspecific similarities in courtship behaviors were measured by calculating Euclidean distances between species based on 12 courtship characters and by clustering species according to UPGMA (unweighted pair-group method using arithmetic averages). The resulting phenogram revealed two major behavioral patterns in courtship: (1) interactive and (2) simple. The former was characterized by a complex sequence in which, typically, a male approached a pheromoneemitting female, engaged in a head- to- head posture with the female, and then brought his abdomen over his head and struck the female on the head and thorax. This action brought male abdominal scent structures into close proximity with the female antennae. The male then attempted copulation from the head- to- head position by a dorsolateral thrust of the abdomen toward the female genitalia. Males of these species possessed scent structures located either on the eighth abdominal segment, or in a costal fold of the forewing, or both. Courtship in the second group was much more prosaic. After locating the female by response to her sex pheromone, the male simply attempted copulation by lateral abdominal thrusts under the female wing, without behavioral embellishments. Males of species exhibiting simple courtship had either no scent structures or structures that appeared vestigial. The grouping of species based on courtship characters was poorly correlated with taxonomic relationships, suggesting that the selective pressures governing the evolution and maintenance of courtship and male pheromones were distinct from those involved in the evolution of other morphological characters. While we argue that the primary force molding the evolution of courtship was an adaptive response to interspecific mating mistakes, we do not believe that isolation is brought about by the sequence of courtship behaviors themselves, due to the striking similarity in the sequence across several diverse species. Rather, these behaviors act to deliver more efficiently the male pheromonal message, which mayhave evolved for reproductive isolation.     

Communication in the courtship of a madagascan hissing cockroach. I. Normal courtship

The hissing Madagascar cockroach, Gromphadorhina portentosa, has a prolonged and complex courtship involving signals in several sensory modalities. Courtship was described for 13 pairs of cockroaches and the frequencies and sequencing of 16 behavioural units were analysed. Particular attention was paid to the function and interactions of acoustic, chemical and tactile components. The results indicate that posturing and sound production by males, and antennation by both males and females, are important in courtship. They also suggest that courtship in G. portentosa, rather than depending on a rigid sequence of behaviour determined by a series of discrete releasers, is quite flexible, using ‘behavioural monologues’ by both sexes as a means of achieving transitions from one stage of courtship to the next.     

Patterns of turning in courtship orientation of the male German cockroach

Courtship turning of the male German cockroach, Blattella germanica, was analysed with respect to rotational and translational movements. Three different sources of orientation information are postulated for the control of this behaviour: external sensory information (perception of sex pheromone) releases courtship, proprioception from the legs is required to curtail turning when the specified turn angle is reached, and internally stored information determines the sign of turning and patterns the courtship actions.     

First observation of the courtship behaviour of the giant devil ray Mobula mobular (Myliobatiformes: Mobulidae)

Courtship behaviour of the giant devil ray Mobula mobular is described from northern New Zealand, temperate southwest Pacific Ocean, for the first time. A mating train consisting of a full-term pregnant female and up to four males was observed over a period of 147 minutes. Their behaviour was similar to courtship behaviour observed in other large mobulids. Biting of the female was not observed, possibly due to the female’s use of the surface to prevent males positioning themselves above her. However, the lead male pressed the female’s abdomen and underside each time the female reached or stopped at the surface. The occurrence of pregnant females and mating behaviour off northern North Island confirms breeding occurs in New Zealand waters.     

Physiological Costs of Repetitive Courtship Displays in Cockroaches Handicap Locomotor Performance

Courtship displays are typically thought to have evolved via female choice, whereby females select mates based on the characteristics of a display that is expected to honestly reflect some aspect of the male’s quality. Honesty is typically enforced by mechanistic costs and constraints that limit the level at which a display can be performed. It is becoming increasingly apparent that these costs may be energetic costs involved in the production of dynamic, often repetitive displays. A female attending to such a display may thus be assessing the physical fitness of a male as an index of his quality. Such assessment would provide information on his current physical quality as well as his ability to carry out other demanding activities, qualities with which a choosy female should want to provision her offspring. In the current study we use courtship interactions in the Cuban burrowing cockroach, Byrsotria fumigata to directly test whether courtship is associated with a signaler’s performance capacity. Males that had produced courtship displays achieved significantly lower speeds and distances in locomotor trials than non-courting control males. We also found that females mated more readily with males that produced a more vigorous display. Thus, males of this species have developed a strategy where they produce a demanding courtship display, while females choose males based on their ability to produce this display. Courtship displays in many taxa often involve dynamic repetitive actions and as such, signals of stamina in courtship may be more widespread than previously thought.     

Functions of courtship feeding in black-legged kittiwakes: natural and sexual selection

Various authors have proposed that courtship feeding evolved under natural selection, sexual selection or both. Using observations of 250 breeding pairs over 3 consecutive years for a total of 77 000 nest-hours, we examined the functional significance of courtship feeding in a seabird, the black-legged kittiwake, Rissa tridactyla. We predicted courtship feeding would benefit males and females in different ways: males may invest in future progeny and females may allow copulations in exchange for food or use courtship feeding to assess males' parental quality. Courtship feeding was correlated with clutch size in 1 of 3 years, suggesting that males may increase their reproductive success by provisioning their mates. Courtship feeding, which was individually repeatable between years, was also related to male arrival date. These results suggest that courtship feeding is a reliable index of male quality that females may use for mate appraisal. Because courtship feeding commences after the pair bond is established, it cannot be used as a criterion for mate choice in the pair's first year. However, courtship feeding rate was related to male re-pairing success in the following breeding season. We thus suggest that courtship feeding in kittiwakes could have first evolved under natural selection (the nutrition hypothesis) and subsequently may have become a signal of male quality that females could use to evaluate their mates for future seasons (future mate appraisal hypothesis). Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Geographic variation in male courtship acoustics and genetic divergence of populations of the Cotesia flavipes species complex

Courtship behaviors of insect populations can vary across the range of a species. Populations exhibiting divergent courtship behavior may indicate genetic divergence or cryptic species. Courtship acoustic signals produced by male wing fanning and genetic structure (using amplified fragment length polymorphisms) were examined for seven allopatric populations of the Cotesia flavipes (Hymenoptera: Braconidae) species complex, using four C. sesamiae (Cameron) and three C. flavipes Cameron populations. Members of this species complex parasitize lepidopteran pests in gramineous crops including sugarcane, maize, and rice . Significant variation was detected in courtship acoustic signals and genetic structure among populations of both species. For C. sesamiae, courtship acoustic signals varied more between populations of two biotypes that were collected near an area of sympatry. The two biotypes of C. sesamiae were also genetically divergent. For C. flavipes, significant differences in acoustic signals and genetic structure occurred among allopatric populations; these differences support the recent designation of one population as a new species. Courtship acoustics play a role in reproductive isolation in this species complex, and are likely used in conjunction with chemical signals. Ecological factors such as host range and host plant use may also influence the divergence of both courtship acoustic signals and genetic structure among populations in the C. flavipes complex.     

Menstrual cycle shifts in attentional bias for courtship language

The current study investigated whether women show an attentional bias toward courtship language during the fertile phase of their menstrual cycle. Thirty heterosexual women (17 naturally cycling, 13 using hormonal contraceptives) completed a dichotic listening task on both a high and low fertility day of their menstrual cycle. Participants were asked to verbally repeat (shadow) an emotionally neutral target passage played in one ear while either a neutral or courtship distracter was played in the other ear. Courtship distracters were flirtatious in content but not overtly sexual. Shadowing errors were coded as a measure of attentional bias toward the distracter. Saliva samples were taken to determine whether levels of estradiol, progesterone and/or testosterone correlated with task performance. As predicted, naturally cycling women made more shadowing errors when listening to a courtship distracter during the fertile phase of their cycle than during the nonfertile phase. This effect was moderated by relationship status, such that fertile, mated women showed an attentional bias for courtship language but fertile single women did not. However, because of small sample sizes in the analysis, this relationship should be viewed as preliminary. Hormonal analysis revealed that higher levels of salivary estradiol predicted greater attentional bias toward courtship language in naturally cycling women. These results suggest that women's attention is drawn to verbal courtship signals when they are fertile, and that this shift is linked to increased estradiol release during the periovulatory phase.     

The extent of visual control in the courtship tracking of D. melanogaster

The courtship tracking behaviour of male D. melanogaster was filmed in a range of visual mutants and in wild type flies under white and infra-red light. The absence of visual input severely restricts the velocity at which the female may be tracked but does not remove the male's ability to discriminate the female's abdomen from her head. Degradation of visual processing, such as occurs in certain visual mutants, leads mainly to disruption of the control of orientation in tracking. The gain control of translation may only be meaningfully assessed when vision is enabled. Courtship tracking does not depend upon the presence of a binocular field of vision; one eyed flies court with a lateral displacement of the body, and loose visual contact with the female readily when her image moves from the sighted towards the blind eye. The selection of the left or right wing for extension follows a distinct pattern with the male's position and orientation around the female; this may be controlled collateral to locomotor output and after interlateral comparison of sensory input.     

Temporal patterning of courtship behaviour in some parasitic Hymenoptera,with special reference to Melittobia acasta

In many parasitic hymenoptera copulation is preceded by elaborate courtship displays which include species-specific characteristics. Other features, shared by related species, may be used for defining higher taxa. The male's repertoire consists of movements involving the wings, legs, antennae, and mouthparts. These movements are performed continuously, or intermittently, depending on the species involved. The elements of a repertoire are repeated over and over again until the female indicates her readiness to copulate. Temporal patterning of various displays, and the timing of the female response are described. In Melittobia acasta (Walker) (Eulophidae) the male display is composed not only of repeating elements, but also includes new elements, introduced along the way; the display progresses towards a climactic finale. The timing of the female's copulation signal is accurately predictable. The morphology of Melittobia males is discussed in relation to this behaviour. Courtship of a related species, M.chalybii, is compared to the courtship of M.acasta.     

The visual control of courtship behaviour in the fly Poecilobothrus nobilitatus

1. Poecilobothrns nobilitatus has a spectacular courtship in which the males perform a complex aerial display directed towards the females. This begins with short bouts of wing-waving, after which the male hovers in front of the female. This is followed by two types of display flight. In one the male circles in an arc around one side of the female, and in the second he flies over her, rotating through 180° in 40ms. The display flights are not modified once they have begun, and are thus pre-coordinated rather than feedback driven, as in tracking behaviour.
2. Courtship behaviour is confined to the males, and the extra tasks the male nervous system is required to perform are discussed in terms of the information needed for the execution of the behaviour.

     

Female goats use courtship display as an honest indicator of male quality

Due to the differential cost of reproduction in promiscuous species, like domesticated goat(s) (Capra hircus), it is expected that females should mate with higher quality males, while males should mate with a greater number of females. Females may use conspicuous secondary sexual characteristics of males such as courtship display to distinguish among high and low quality males. Testosterone (T) controls a large suite of secondary sexual characteristics and variation in T concentrations may account of differences in courtship rates. Two studies were conducted to examine the relationship between T concentrations and courtship rate and its role in mammalian female mate choice. Experiment 1 utilized bucks (intact males) and Experiment 2 utilized T-replaced wethers (castrated pre-pubertally). During the first year of Experiment 2, T-replaced wethers received either vehicle control (CON), 25 mg or 100 mg testosterone propionate (TP). During the second year of Experiment 2, T-replaced wethers were treated with either 5 mg, 15 mg or 25 mg TP. For all experiments, mean courtship rates and circulating T concentrations were measured, as well as female preference for males displaying different courtship frequencies. T concentrations and courtship rate were positively correlated for bucks and estrous females preferred high courting bucks. Males receiving 25 mg and 100 mg TP courted females at a similar rate, but both were significantly higher than courtship rates of the CON wethers. Courtship rates of the 25 mg, 15 mg and 5 mg TP-treated males were all significantly different. Females did not show a preference between the 100 mg or 25 mg TP-treated wethers, however both were preferred in comparison to the CON wethers. Females did not show preference between the 25 mg or 15 mg TP-treated wethers, however both were preferred in comparison to the 5 mg TP-treated wethers. Taken together, these studies suggest that courtship rate is T-dependent. Further, females can use courtship rate to distinguish among males.     

Excitatory and inhibitory switches for courtship in the brain of Drosophila melanogaster

BACKGROUND: Courtship is the best-studied behavior in Drosophila melanogaster, and work on its anatomical basis has concentrated mainly on the functional identification of sexually dimorphic sites in the brain. Much less is known of the more expansive, nondimorphic, but nonetheless essential, neural elements subserving male courtship behavior. RESULTS: Sites in the CNS mediating initiation and early steps of male courtship in Drosophila melanogaster were identified by analyzing the behavior of mosaic flies expressing transgenes designed either to suppress neurotransmission or enhance neuronal excitability. Suppression of neurotransmission was accomplished by means of the dominantly acting, temperature-sensitive dynamin mutation shibire(ts1), whereas enhanced neuronal excitability was produced by means of a novel, dominantly acting, truncated eag potassium channel. By using a new, landmark-based procedure for aligning diverse expression patterns among the various mosaic strains, a comparison of courtship performance and affected brain sites in strains expressing the transgenes identified a cluster of cells in the posterior lateral protocerebrum that exerts reciprocal effects on the initiation of courtship, suppressing it when they are inactivated and enhancing it when they are hyperactivated, indicative of cells that normally play an excitatory, triggering role. A separate group of nearby cells, slightly more anterior in the lateral protocerebrum, was found to inhibit courtship when its activity is enhanced, indicative of an inhibitory role in courtship. CONCLUSIONS: A cluster of cells, some excitatory and some inhibitory, in the lateral protocerebrum regulates courtship initiation in Drosophila. These cells are likely to be an integration center for the multiple sensory inputs that trigger male courtship.