Seasonal variation in mate-locating activity in the desert hackberry butterfly (Asterocampa leilia; Lepidoptera: Nymphalidae)

Males of the desert hackberry butterfly occupy and defend encounter sites used in mate location. This activity is restricted to about 3 h each day, usually between 0800 and 1100. We monitored daily and seasonal changes in the number of males perched at encounter sites along a census route on 32 days during 1991 and 1992. The results indicate that males are active earlier in the mornings during summer than in spring or fall. However, they were not active at all times when air temperature and solar altitude would appear to permit perch site occupation. Daily and seasonal shifts in air temperature and solar radiation are not a sufficient proximate explanation for patterns of male behavior. Over a wide range of temperatures, females most often eclose around 0800, which suggests that the timing of female availability determines when males are active within the broad limits imposed by thermoregulatory requirements.     

Copulatory courtship and cryptic female choice in red flour beetles Tribolium castaneum

Males of many animal species engage in courtship behaviours during and after copulation that appear to be solely aimed at stimulating the female. It has been suggested that these behaviours have evolved by cryptic female choice, whereby females are thought to impose biases on male postmating paternity success. Males of the red flour beetle Tribolium castaneum rub the lateral edges of the females' elytra with their tarsi during copulation. We manipulated female perception of this behaviour by tarsal ablation in males, thus preventing males from reaching the edge of the female elytra with their manipulated legs, and by subsequently performing a series of double-mating experiments where the copulatory behaviour was quantified. We found a positive relationship between the intensity of the copulatory courtship behaviour and relative fertilization success among unmanipulated males. This pattern, however, was absent in manipulated males, where female perception of male behaviour differed from that actually performed. Thus, female perception of male copulatory courtship behaviour, rather than male behaviour per se, apparently governs the fate of sperm competing over fertilizations within the female, showing that copulatory courtship is under selection by cryptic female choice.     

Male behaviour and female choice: Mate selection in the wren?

The European wren is frequently polygynous. Males occupy exclusive territories within which they spend much time displaying and building nests. They lead females to their nests in courtship. Males building the most nests during a season also make the most breeding attempts. Data collected in one of two years suggested that individual females may show a preference for males with the largest number of complete but vacant nests. There was no consistent correlation between male display behaviour and female mate choice. Data on breeding success may suggest why polygyny is adaptive for both males and females in this species.     

Mating in the mosquito, Anopheles gambiae s.l.

ABSTRACT. Swarming behaviour in the Anopheles gambiae complex was observed in the field, in the Gambia, West Africa, and in the laboratory. Naturally occurring swarms of A.melas were seen in a clearing at the edge of mangrove swamps close to their breeding sites. Males could be induced to swarm over artificial 'markers' within this 'arena' but not outside it. Females were observed entering the swarm and mating. In the laboratory, in an artificial 'dusk', male A.gambiae s.str. swarmed over a black marker on the floor of their 1.2-m cube cage. In contrast to the males, females made only short flights over the marker, performing brief turning movements at its edge. It is proposed that swarming brings about the aggregation necessary before short-range attraction can take place, and that, in nature, anopheline mosquitoes orientate visually first to an arena and then to a marker within the arena. Female behaviour can be interpreted as a process of scanning possible swarm sites until mating is achieved.     

Female choice of sites versus mates in a coral reef fish,Thalassoma bifasciatum

To understand the dynamics of sexual selection, one needs to identify the basis of female choice in mating. Daily censuses showed that most spawns of individual female bluehead wrasses, Thalassoma bifasciatum, occurred at one of several available mating sites, either in a group-spawning aggregation of small males or with a large territorial male. Most females were not located near their principal mating site during the non-spawning period of the day. Females were thus exposed to other males and sites and should have had the opportunity for choice. When a resident territorial male disappeared or was removed experimentally from a mating site, females did not alter their fidelity to that site. Furthermore, manipulations showed that when a territorial male changed the location of his mating activity to an adjacent site, females did not follow him but continued to mate at their customary site. Thus mating sites, rather than the males occupying them, appeared to be the objects of female choice. The bright coloration and courtship behaviour of larger males may demonstrate the safety of a particular site rather than indicating some aspect of male quality.     

Elaborate Nests in a Weaverbird: A Role for Female Choice?

Studies on sexual selection have focused on behaviour and morphology, but several groups of animals build elaborate structures associated with acquiring a mate. I investigated female choice for nests built by male baya weavers (Ploceus philippinus). Nest choice by females should be strong, as nests are obvious direct benefits provided by males. I used a field experiment supplemented with correlational information to ask whether females appear to base mate choice decisions on male behaviour, nest architecture, and nest location. When the nests of highly visited males were exchanged with those of poorly visited males, female visits remained highest at the original male and location. I found no relationship between female choice and male display or other behaviour. Correlational analyses show that nest location was a better predictor of female choice than was nest architecture. These data suggest that current female choice is driven more by access to safe nesting sites rather than to well‐built nests, possibly because all males are able to build nests of adequate quality. However, nest architecture is unlikely to be irrelevant to females, and its role deserves further investigation.     

Male dwarf chameleons assess risk of courting large, aggressive females

Conflict between the sexes has traditionally been studied in terms of costs of mating to females and female resistance. However, courting can also be costly to males, especially when females are larger and aggressively resist copulation attempts. We examined male display intensity towards females in the Cape dwarf chameleon, Bradypodion pumilum, in which females are larger than males and very aggressive. We assessed whether aggressive female rejection imposes potential costs on males and whether males vary their display behaviour with intensity of female rejection, female size or relative size differences. Males persisted in courtship after initial female rejection in 84% of trials, and were bitten in 28% of trials. Attempted mounts were positively associated with males being bitten. Males reduced courtship with increased intensity of female rejection. Male courtship behaviour also varied with female size: males were more likely to court and approach smaller females, consistent with the hypothesis that larger females can inflict more damage. These results suggest that, in addition to assessing female willingness to mate, male dwarf chameleons may use courtship displays to assess potential costs of persistence, including costs associated with aggressive female rejection, weighed against potential reproductive pay-offs associated with forced copulation.     

Feather stealing in the satin bowerbird (Ptilonorhynchus violaceus): male competition and the quality of display

Male satin bowerbirds use feathers to decorate their bowers and often steal feathers and other decorations from the bowers of other males. Decorations are a key element in sexual display and tracking their movement between bowers provides the first detailed information about this unique pattern of sexual competition. For two field seasons the movement of marked feathers was followed. Males varied greatly in stealing activity. The most active feather thieves were often from areas where bowers were close together and they were involved in reciprocal stealing with males at adjacent bowers. The rate of stealing by males was significantly correlated with the number of feathers on their bowers. This suggests that stealing is important in determining the level of bower decoration and mating success. Patterns of stealing behaviour support models of sexual selection which suggest that male interactions are important in influencing female choice through their effect on the quality of male display.     

Male perch selection and the mating system of the robber fly,Promachus albifacies (Diptera: Asilidae)

We studied the activity and spatial distribution of the robber fly,Promachus albifacies, in a desert grassland habitat in central New Mexico. Late in the season males spent most of the daytime on or near cholla and yucca plants that had dead stems or dead flower stalks at least 1 m high. Of the three hypotheses (thermoregulation, foraging, mate encounter site) considered as explanations for this distribution, the mate-encounter-site hypothesis was best supported. Plants used by females as oviposition sites were the focus of male activity. Males perched within or near these plants and attempted copulations with females detected nearby. Most matings were initiated at these locations. Seasonal changes in male and female activity also supported the mate-encounter-site hypothesis. Early in the season, females spent little time ovipositing, and predictably, males spent little time on or near these plants. Such a mating system may be described as resource defense polygyny, since males acted aggressively toward one another at oviposition sites even when females were not present. However, the short tenure of males at these sites is suggestive of scramble competition polygyny. We discuss possible reasons why this particular mating system has evolved.     

Sexual Differences in the Responses of Captive Great Reed Warblers (Acrocephalus arundinaceus) to Variation in Song Structure and Repertoire Size

The function and evolution of repertoires in songbirds is still not clearly understood. This study involves the great reed warbler Acrocephalus arundinaceus and playback of recorded song to both captive males and females under controlled laboratory conditions. Females were implanted with oestradiol and the copulation solicitation display used as an index of sexual response. Males were not implanted and crest-raising used as an index of aggressive response. Two types of songs were used (short and long) and two sizes of repertoire (high and low). Females only responded to long songs, and responded significantly more to high than low repertoires. It was concluded that long songs have a sexual function and that high repertoires are the result of intersexual selection through female choice. Males responded to short and long songs, but did not discriminate between high and low repertoires. Adding a live male to the experiment made no difference to female responses but males responded significantly more. When field studies are also taken into consideration the overall conclusion is that song quality and repertoire size is particularly important in female choice, and that short songs and a male presence are more important in aggressive behaviour. Integration of both field and laboratory studies on males and females should lead to more rigorous testing of hypotheses concerning the function and evolution of bird songs.     

Preliminary study on female choice of mates versus sites in a wrasse, Cirrhilabrus temminckii

Female choice of mates versus sites was studied in a wrasse, Cirrhilabrus temminckii. Males had territories within a restricted area on a rocky slope at which females visited and pair-spawned pelagic gametes. Females visited several males or territories before spawning, suggesting the opportunity of female choice. Of the four characteristics of territorial males examined—body size, ratio of pelvic fin length to body size, courtship, frequency and territory depth—only territory depth was significantly correlated with daily mating success of males. The former three male characters were not related to territory depth. These results suggest that female C. temminckii chooses deep sites rather than specific mates in mating.     

Alternative mating strategies in a desert grasshopper: evidence of density-dependence

Field observations revealed that, on a given day, male mating behaviour in a population of the grasshopper Ligurotettix coquilletti ranged from little or no stridulation (inactive) to relatively persistent singing (active signalling). Inactive males were usually located in the territories of active signallers. Actively signalling male achieved more matings, and also more frequently approached and mounted females in incidents terminating in the male departing the female without copulating. Individuals switched between inactive and actively-signalling behaviour during their adult lives, and males that were usually active signallers achieved greater lifetime mating success. Thus, we obtained no evidence to support the hypothesis that variation in mating behaviour was maintained by negative frequency-dependent selection. By comparing the behaviour of males in two field plots maintained at different population densities, however, we found that high density was directly related to a higher incidence of inactive behaviour. Males perched on creosote (Larrea tridentata) bushes (their host plant), and certain bushes. Mating behaviour was independent of body size and, while young (<7 days old) males tended to be inactive, neither age nor the time of adult maturation during the season could fully account for the incidence of inactive behaviour among males. Instead, we suggest that the adoption of inactive behaviour resulted partly from aggressive encounters between males.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.     

Are Centrally Displaying Males Always the Centre of Female Attention? Acoustic Display Position and Female Choice in a Lek Mating Subterranean Insect

Several conceptual models seek to explain patterns of male display and factors that influence female mate choice in lek mating systems. The central advantage model predicts that males displaying at or near the lek centre should be more attractive to females than are males positioned along the lek periphery. Females may exhibit biases toward these centrally displaying males based on either spatial or display‐related cues. We tested the prediction of the central advantage model in investigating the importance of male display position in the subterranean and lek mating prairie mole cricket (Gryllotalpa major). Gryllotalpa major males form mating aggregations in the early spring and produce an acoustic advertisement signal from a constructed calling chamber at the soil surface. Pair formation occurs in the calling chamber, and males typically maintain these structures for the duration of the reproductive season. To assess whether G. major females exhibit a preference for males calling from centrally located acoustic burrows, we documented the spatial position and number of female attractions for all advertising males across the focal lek. Six spatial attributes related to display position were reduced using principal component analysis and examined for an association with male attractiveness. We found that in general, female attractions were distributed randomly across the lek; male attractiveness was not related to proximity to the lek centre nor to any factor associated with display position. The most highly attractive males, however, were located further from the lek centre and from nearest calling neighbours than other attractive males. Advertising males that segregate themselves within the aggregation and locate nearer the lek margin may gain a geometric advantage resulting in the increased probability of attracting a searching female.     

Cross-fostering Does Not Influence the Mate Preferences and Territorial Behaviour of Males in Lake Victoria Cichlids

The ability to recognise conspecifics in contexts of mate choice and territorial defence may have large effects on an individual's fitness. Understanding the development of assortative behaviour may shed light on how species assortative behaviour evolves and how it may influence reproductive isolation. This is the case not only for female mate preferences, but also for male mate preferences and male territorial behaviour. Here we test with a cross-fostering experiment whether early learning influences male mate preferences and male–male aggression biases in two closely related, sympatrically occurring cichlid species Pundamilia pundamilia and Pundamilia nyererei from Lake Victoria. Males that had been fostered, either by a conspecific female or by a heterospecific female, were tested for their aggression bias, as well as for their mate preferences, in two-way choice tests. Males cross-fostered with conspecific and heterospecific foster mothers selectively directed their aggression towards conspecific intruders. The cross-fostering treatment also did not affect male mate preferences. These results are in striking contrast with the finding that females of these species show a sexual preference for males of the foster species.     

Choice of Lookout Posts by Territorial Amberwing Dragonflies, Perithemis tenera (Anisoptera: Libellulidae)

Many territorial insects use specific perches, or lookout posts, from which they perceive and react to conspecific males and females. We investigated the lookout post choice of territorial male amberwing dragonflies (Perithemis tenera). An observational study indicated that males rarely perched directly at their oviposition site; rather, they perched farther out from shore than, and within 2 m from, their oviposition site. In an experimental study, we provided an array of perches at different distances from the shore and oviposition site to eliminate perch limitation as a factor in perch choice. The results of the experimental study confirmed the patterns evident in the observational study; males perched farther from shore than their oviposition site was located. Interestingly, in both the observational and the experimental study, when neighbors were close, a male perched away from his closest neighbor, which usually resulted in his oviposition site being closer to his neighbor than he was. Thus, male amberwings apparently alter their perch choice within their territories in response to the location of other males. These lookout post locations may provide the best opportunity for the territorial male to perceive passing females and intruding males, while minimizing conflict with their neighbors.     

Differences in size and energy content affect the territorial status and mating success of a neotropical dragonfly

In male odonates, both size and fat content are related to territory defence and mating success. Males that are larger and have higher energy reserves win relatively more disputes for territory and attract more females. Wing colour has also been regarded as a mechanism that influences agonistic behaviour between males, as wing pigmentation might be regarded as a sign of male quality. In this study, we analysed whether a set of male physical (body size and wing colour), physiological (body fat content) and behavioural (disputes between males) characteristics were involved in the territory defence and mating behaviour of the neotropical dragonfly Zenithoptera lanei Santos, 1941 (Anisoptera: Libellulidae). Males were characterised as territorial whenever they warded‐off other males and remained in the same place within the pond for two consecutive days. In general, these territorial males were larger and had more abdominal and thoracic fat, engaged in pursuits more frequently, spent more time on sexual behaviour and female guarding, and mated more in comparison to subordinate males. By evaluating whether the percentage of wing area covered by black ink influenced male behaviour, we found that territorial males tended to act aggressively towards other males whose wings were partially painted, and sexually towards females irrespective of wing area painted. In Z. lanei, both body size and fat content play a role in defining territoriality. By subduing competitors and dominating preferred locations within high‐quality sites, these males are likely to be visited by females and engage in mating.     

Male territoriality and mating system in the European beewolf Philanthus triangulum F. (Hymenoptera: Crabronidae): evidence for a “hotspot” lek polygyny

The evolution of the mating system of a species is strongly influenced by the spatial and temporal distribution of females and/or resources. Here, we describe aspects of the territorial behavior of males of a solitary digger wasp, the European beewolf (Philanthus triangulum) and characterize the mating system. We show that beewolf males establish small territories that do not contain any resources essential to females. These territories are intensively scent-marked with a pheromone from a cephalic gland and are defended against intruders in combat flights. We provide evidence that scent-marking constitutes a chemical display and that the pheromone serves to attract receptive females and, thus, represents a sex pheromone. Using spatial statistics, we show that beewolf territories are clumped in space both with respect to other male territories and, more importantly, with respect to female nesting sites. Additionally, the proportion of days a territory is occupied by a male is correlated with the number of female nests in the vicinity. Taking into account that beewolf males do not defend or provide resources essential to females, but merely display chemically to attract females for mating in an aggregation of territories close to female nesting sites, we conclude that the European beewolf exhibits a hotspot lek polygyny with female nesting sites constituting “hotspots” for lek formation.     

Big is better: the importance of size for reproductive success in male Pomatoschistus minutus (Pallas) (Pisces, Gobiidae)

The sand goby, Pomatoschistus minutus , was used in aquarium experiments to study the importance of body size for the ability of males to gain access to nest sites and mates. When several male and female P. minutus were allowed to reproduce together, on average, half of the males built nests, and half of those males obtained eggs in their nests. Males with nests were significantly larger than males without nests, and nests with eggs belonged to males larger than the males with empty nests. In another experiment, when two males were competing for one nest, the largest male occupied the nest when both fish were put into the aquarium simultaneously. However, when the smaller male had been allowed to establish a nest before the larger male was introduced, the small male could usually retain the nest. Males with a body length < 50 mm did not build nests at all in the early part of the breeding season. In female choice experiments, no preference for larger males was found. Thus, male-male competition for nest sites and behavioural differences between different sized males seem to be the main factors influencing the non-random mating success in male P. minutus.