Characterisation of Predator‐Directed Displays in Tephritid Flies

Flies of the family Tephritidae are known to perform a display in front of their salticid spider predators. This display involves extending the wings while the fly moves from side to side. The wings of many of fly species are banded, and in some species, these bands are thought to deter their predators by mimicking the leg patterns of salticids. However, as this display is also seen in non‐mimicking flies, there is some uncertainty over the functional significance of these displays. In this study, we explored the efficacy of displays by the lightly banded but non‐mimicking Mexican fruit fly (Anastrepha ludens) in deterring attacks by the salticids Paraphidippus aurantius and Phidippus bidentatus. We tested the effect of band visibility and the fly's physiological condition on the production of these displays and the probability of attack by salticids. We filmed the interactions of flies and spiders and analysed fly displays in the presence of the salticid. We show that the fly's display deters salticids and promotes the fly's chances of survival and that the presence or absence of bands does not alter the possibility of attack. Flies fed on different quality diets showed similar rates of display. Our results suggest that tephritid flies deter attacks because of their display and not their appearance.     

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.     

Effects of visitors and cage changes on the behaviors of mangabeys

Three groups of golden-bellied mangabeys were studied to determine the effects of visitors and cage changes on their aggressive displays, grooming and sexual behavior, and play. Assessments of changes in behavior were made by comparing categorized observer comments. Animals moved to cages having more visitors increased their aggressive displays toward people, decreased their aggressive displays toward other species in neighboring cages, and increased their withingroup aggression. On the other hand, grooming, sexual behavior, and play increased after cage changes regardless of numbers of visitors. The view that it is the pattern of behavior changes in enrichment that is important in assessment of attempted enrichment was supported. Aggressive and affiliative behaviors are affected quite differently by different environmental stimuli in the zoo.     

The function of male aggressive displays towards females in mountain gorillas

In groups ofGorilla g. beringei, male aggression towards females regularly takes the form of male display. This paper examines male display towards females in two Karisoke study groups (Group 5 and Group BM) in 1989, a period when none of the females were new immigrants. Results are based on 259 hr of focal observations and 121 hr ofad libitum observations on male behaviors towards females. The goal is to see if the data are compatible with four non-mutually exclusive hypotheses to explain male displays towards females: (1) demonstration of male fighting abilities to influence female long term residence decision; (2) decrease potential competitive inequities between females; (3) provision to females of an occasion to confirm their subordinance to a male; and (4) short term influence on mating. First, male-female proximity was tested against proportion of male displays, to rule out the possibility that males display towards females simply because they happen to be close by. There was no association between proximity and male display. Dominant males were responsible for a higher proportion of displays than subordinate males. This is consistent with the idea that males display to demonstrate their fighting abilities, or their qualities as protector, since dominant males are the ones offering long term protection against infanticide and predators. Females that were in a position to transfer did not receive a higher proportion of male display, however. Long term resident dominant females received a higher proportion of displays from the dominant males, which is consistent with the idea that males attempt to decrease potential competitive inequities between females. There was an association between female appeasement reactions and male displays, which suggests that males display to create occasions for the females to confirm their subordinance to them. Estrous females did not receive a higher proportion of male displays, and there was no association between male display and copulation, suggesting that male displays are not a form of courtship aggression aimed at influencing mating in the short term.     

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 Function of Male Agonistic Displays in Ursine Colobus Monkeys (Colobus vellerosus): Male Competition,Female Mate Choice or Sexual Coercion?

Male agonistic displays may allow males to assess competitors, females to assess mates, or could be directed at cycling females to sexually coerce them. We analysed the display output of 26 male ursine colobus monkeys (Colobus vellerosus) in four groups over 13‐mo at the Boabeng‐Fiema Monkey Sanctuary, Ghana. Display indices (including three behaviours, loud calls, stiff‐legs, and jump‐displays) were calculated for males in each group. Males vary in their expression of these behaviours suggesting they are sexually selected signals. We investigated the target of displays and whether display indices varied in relation to male dominance rank, eviction of other males, copulation rate, and proceptive behaviours received from females, to assess the primary function of these behaviours. Male displays decreased in vigour over time and were targeted to other groups and males. High‐ranking males displayed more than low‐ranking males. Alpha male display indices correlated with the number of other males evicted from the group. Display rates were generally higher when cycling females were present in the group. However, neither male display index nor rank correlated with copulation rates. Alpha and non‐alpha males gave cycling females equal rates of sexual solicitations; likewise cycling females showed no difference in the rates of proceptive behaviours directed towards alpha and non‐alpha males. Females mated promiscuously and did not seem to base mating decisions on male display output, although data on female hormones is needed to determine if they mate with strongly displaying males more in the periovulatory period. The male–male competition hypothesis received the greatest support, with some support for the female mate choice hypothesis. Although behaviours that appeared sexually coercive were observed, the function of male displays did not seem to be sexual coercion. Displays were rarely directed at females and males that displayed more did not have greater mating success.     

Male Golden‐collared Manakins Manacus vitellinus do not adapt their courtship display to spatial alteration of their court

Male Golden‐collared Manakins Manacus vitellinus perform an elaborate courtship display composed of acrobatic jumps between saplings delimiting a court on the forest floor. Males rehearse their displays for hours until they are executed with amazing precision and speed. Here we investigated the plasticity of the display by examining whether males modify their choreography in response to a disturbance, such as when a small branch is placed against one of the saplings. Male Manakins performed displays that were longer and lacked a key element that invites females to copulate, supporting the hypothesis that males learn a specific sequence of moves to build their choreography.     

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Courtship displays are typically comprised of the same behavioral pattern, or patterns, repeated several times by males. Both the quantity and quality of the displays produced by a given male bird are not, however, constant. The number and/or quality of displays can decrease over time, indicating fatigue, or males can increase the number and/or quality as they display more, indicating a warm-up period. Although there is evidence for fatigue or warm-up periods for many types of courtship displays, data on motor components of avian courtship are scant, despite how commonly they are used. Here, we test whether drumming, a non-vocal motor display, in male ruffed grouse (Bonasa umbellus) changes in relation to the number of displays executed. Using a large number of recordings, our linear mixed models yielded a significant effect of cumulative number of drumming displays on the number of wingbeats per second, referred to as pulse rate. Across males, pulse rate is slowest when males begin drumming each day and increases until approximately 50 drumming displays have been produced. The rate of increase is also modulated by the nighttime low temperature such that cooler conditions are associated with lower pulse rates and a slower increase in pulse rate relative to the cumulative number of displays. Further, the maximum pulse rate recorded and average pulse rate after 50 displays is inversely correlated with body mass such that larger males are slower than smaller males. We suggest that the daily changes in pulse rate likely reflect a warm-up period based upon the effects of cumulative drumming count and temperature on pulse rate. Whether these dynamic changes in the production of a motor display are informative to female grouse is unknown. However, we propose that daily changes in how motor displays are performed may be a common feature of avian courtship that has gone relatively unnoticed, despite the potential for motor performance to be a trait that is important for female mate choice.     

Sizing‐up the Competition: Factors Modulating Male Display Behavior During Mate Competition

When competing for mates, males of many species use cues from their rivals to evaluate their chances of success. Signaling behavior is a vital component of male–male contests and courtship, and may inform males of a rival's quality or intentions. We used eastern fence lizards (Sceloporus undulatus) to investigate how the time a male spent signaling during mate competition is influenced by his quality, his rival's quality relative to his own, and the value of a contested female. Furthermore, we examined how a male's behavioral response to a competitor's signals would be mediated by his relative quality. We simulated natural encounters by allowing two males to compete over a single female in the laboratory. We measured the time males spent performing two types of displays (pushups and shudders) and categorized male behavioral responses to rival pushup and shudder displays. Time spent signaling was not related to a male's absolute quality (body and head size, condition, and badge sizes), or his quality relative to that of his rival, although males did spend more time performing pushups when competing over females in better condition. Male behavior was also influenced by his rival's signals, such that males of relatively lower quality than their opponents were more likely to aggressively respond to rival pushups and shudders. We discuss these results with respect to the evolution and function of signaling behavior in courtship and male–male contests.     

Sex differences in antipredator tail-waving displays of the diurnal yellow-headed gecko Gonatodes albogularis from tropical forests of Colombia

Many vertebrate species show display behaviors when predators are in their vicinity. Some of these displays may inform the predator of the improbability of capturing the prey (i.e., pursuit-deterrent displays) and are potentially advantageous to both predator and prey. Here we present data on a tail display performed by Gonatodes albogularis, a diurnal tropical gecko. We performed transect surveys in three habitats near Bogotá in Colombia. Geckos detected during transects were approached by the observer in a standardized way, and details of their tail-waving displays were recorded. In control recordings animals were watched from a distant site without approaching them. Results showed sexual differences in tail-waving display: when approached by the observer, males performed this behavior more frequently than females. We found no significant differences between males and females in flight-initiation distances and height above the substratum when they were initially located. Results also showed that males displayed more frequently when approached than when the simulated predator remained stationary. We interpret these results as evidence that the display functions as a pursuit-deterrent signal to potential predators. However, as some tail displays were performed in the presence of conspecifics, the display may also have a social function.     

Fragrances, male display and mating behaviour of Euglossa hemichlora: a flight cage experiment

Abstract. Male euglossine bees (Apidae: Euglossini) collect volatile substances (fragrances) from floral and nonfloral sources and store them in hair‐filled cavities in their hind tibiae. Over time, males accumulate large quantities of complex and species‐specific blends of fragrances. Various hypotheses have been put forward to explain this behaviour, including the idea that fragrance stores reflect the genetic quality of individual males and have evolved through sexual selection and female choice. Clear support of this hypothesis is lacking, largely because male–female interactions are both rare and difficult to observe in nature. Here, we report a flight cage experiment performed in Panama that permitted mating between virgin females (raised from brood cells) and males captured in the forest at fragrance baits. In the cage, eight individually marked males defended small territories around vertical perch sites and showed a characteristic display, which included a previously unreported ‘leg‐crossing’ movement, possibly related to fragrance release. A total of six copulations and three copulatory attempts by Euglossa hemichlora were observed and partly recorded on video. The copulations, all of which were initiated by the female landing on a male perch, were short (4–10 s) and showed no signs of the transfer of chemical substances from male to female. In some cases, the male hovered directly over the female before descending to mount her, possibly facilitating fragrance evaluation by the female. After the experiment, the contents of the males' hind legs were analysed by gas chromatography‐mass spectroscopy, which detected complex mixtures of terpenoids and aromatics (totalling 70 different compounds) dominated by hexahydrofarnesyl acetone, farnesene epoxide, ocimene and p‐dimethoxy benzene. Individual total amounts of fragrances were neither related to display activity or perch occupancy by given males, nor to the frequency of matings achieved. Display activity was the only positive correlate of mating frequency. Generally, individuals had uniformly large amounts of stored fragrances in comparison to a previous study of three other species of Panamanian Euglossa.     

Prominent ornaments and rapid color change: use of horns as a social and reproductive signal in unicornfish (Acanthuridae: <Emphasis Type="Italic">Naso</Emphasis>)

The behavior of two unicornfishes, Naso unicornis with a horn-shaped protuberance on the forehead and Naso vlamingii with a round protuberance, was observed in social and reproductive contexts in an aquarium. Males of both species performed displays that were associated with quick changes in the colors of the protuberance and other body parts, highlighting the protuberance by color contrast. The displays with color changes of the protuberance took place when a male courted a female in the evening or in the night. The same displays were occasionally performed by males toward females throughout the daylight hours. In N. vlamingii, dominant males displayed the protuberance toward subordinate smaller males. Although the sizes and shapes of the protuberance were sexually monomorphic, females of both species rarely made displays of the protuberance. A hypothesis about the function of the protuberance in unicornfishes is proposed: that males use the protuberance as a conspicuous signal in courtship and contests among males by emphasizing it with quick changes of its color contrasts. We argue that the combination of morphologically distinct protuberances and quick changes of their color is a sexually selected trait among unicornfishes, because these characters play important roles in intersexual and intrasexual interactions.     

Mating behavior ofPhytoecia rufiventris Gautier (Coleoptera: Cerambycidae)

Sexual behavior between males and females, as well as between males, is described and discussed for the cerambycid beetlePhytoecia rufiventris. The beetles' taxis toward plants taller than average height brings the sexes together from a distance. A male may mount another individual (male or female) and attempt copulation without sex discrimination. The male can discern the sex of another individual only when the terminal part of his abdomen touches the ventral surface of the fifth visible sternite of the latter. No evidence of a sex pheromone is found in this species. Within 1.5–5.5 cm the substrateborne vibrations produced by a moving individual may be the important factor which elicits males to approach a moving individual and attempt copulation. If a female is receptive when a male touches her, he can copulate with her without any courtship display. However, if the female runs away and appears unreceptive, the male will perform courtship displays. Copulation is usually terminated by males. Homosexual behavior between males is discussed.     

Salamander territoriality: pheromonal markers as advertisement by males

A laboratory experiment was performed to test the hypothesis that male red-backed salamanders (Plethodon cinereus) use pheromones contained in faecal pellets to identify male-marked territories. Each of 25 males was tested randomly under four conditions: (1) a burrow marked with its own faecal pellet versus one marked with a conspecific's pellet; (2) own-marked versus surrogate-marked burrows (the surrogate being a pellet of wadded paper); (3) conspecific-marked versus surrogate-marked burrows; and (4) a control of two surrogate-marked burrows. Males spent significantly more time in own-marked than in conspecific-marked burrows and significantly more time in surrogate-marked than in conspecific-marked burrows. Males favoured own-marked over surrogate-marked burrows. No position bias was found in the control. Males spent significantly more time nose-tapping (olfactory sampling) to a conspecific's pellet when it was paired with a surrogate but showed no differences in the other three tests. Significantly more time was spent in a submissive posture in front of the conspecific-marked burrow than in front of either their own-marked or the surrogate-marked burrows; no difference was found between own-marked versus surrogate-marked burrows or in the control. Time spent in the threat posture did not differ significantly between burrows in any condition. These data permit the inferences that males of P. cinereus use faecal pellets to mark and identify territories, avoid or display submissively toward burrows marked by conspecific males, and prefer own-marked shelters.     

Hind Wing Shape Evolves Faster than Front Wing Shape in Calopteryx Damselflies

Wing shape has been shown in a variety of species to be influenced by natural and sexual selection. In damselflies, front- and hind wings can beat independently, and functional differentiation may occur. Males of Calopteryx damselflies show species-specific nuptial flights that differ in colour signalling with the hind wings. Therefore, hind wing shape and colour may evolve in concert to improve colour display, independent of the front wings. We predicted that male hind wing shape evolves faster than front wing shape, due to sexual selection. Females do not engage in sexual displays, so we predicted that females do not show differences in divergence between front- and hind wing shape. We analysed the non-allometric component of wing shape of five European Calopteryx taxa using geometric morphometrics. We found a higher evolutionary divergence of hind wing shape in both sexes. Indeed, we found no significant differences in rate of evolution between the sexes, despite clear sex-specific differences in wing shape. We suggest that evolution of hind wing shape in males is accelerated by sexual selection on pre-copulatory displays and that this acceleration is reflected in females due to genetic correlations that somehow link the rates of wing shape evolution in the two sexes, but not the wing shapes themselves.     

Courtship dives of Anna's hummingbird offer insights into flight performance limits

Behavioural displays are a common feature of animal courtship. Just as female preferences can generate exaggerated male ornaments, female preferences for dynamic behaviours may cause males to perform courtship displays near intrinsic performance limits. I provide an example of an extreme display, the courtship dive of Anna''s hummingbird (Calypte anna). Diving male Anna''s hummingbirds were filmed with a combination of high-speed and conventional video cameras. After powering the initial stage of the dive by flapping, males folded their wings by their sides, at which point they reached an average maximum velocity of 385 body lengths s⁻¹ (27.3 m s⁻¹). This is the highest known length-specific velocity attained by any vertebrate. This velocity suggests their body drag coefficient is less than 0.3. They then spread their wings to pull up, and experienced centripetal accelerations nearly nine times greater than gravitational acceleration. This acceleration is the highest reported for any vertebrate undergoing a voluntary aerial manoeuvre, except jet fighter pilots. Stereotyped courtship behaviours offer several advantages for the study of extreme locomotor performance, and can be assessed in a natural context.     

Homosexual Mating Displays in Penguins

More than 50 yr ago, field studies recorded the same‐sex pairs (and trios) of penguins displaying to each other during the mating season, using behavior patterns typical of heterosexual mating displays. Such observations led to a hypothesis that due to a lack of sex recognition pairing occurs at random with respect to sex, an idea countered by the argument that sex recognition is highly accurate. No quantification of same‐sex mating displays has tested the frequency of such displays in penguins or tested the hypothesis of random display partners with respect to sex. During their mating season, we studied displaying and paired king penguins, Apenodytes patagonicus, at Kerguelen Island and sexed them using a DNA marker, to quantify any occurrence of this behavior. Indeed, same‐sex courtship displays were common (28.3% of 53 displaying pairs), the great majority of which were between males. Some homosexually displaying males eventually paired with females, but such males were significantly slower in heterosexual pairing than males that did not display homosexually. In two extraordinary cases, same‐sex pairs learned each other’s calls, an essential step in the pairing process. The frequency of such pairs was much lower than among displaying couples, significantly so for males. Finally, the frequency of homosexually displaying pairs was significantly lower than expected from random assortment of displaying birds, for both males and females. We examined possible explanations for same‐sex display and its biological significance. A population sex‐ratio bias in favor of males and high concentration of male sex hormones may help to explain non‐reproductive homosexually displaying pairs.     

Harderian letdown in male mongolian gerbils (Meriones unguiculatus) contributes to proceptive behavior

Adult female gerbils in estrus, like other female rodents, tend to engage in proceptive displays toward conspecific males. The displays, which may be interspersed with the more usual female agonistic activity, if the males are strangers, are preceded by female olfactory investigation of those head areas in the male gerbil where Harderian letdown accumulates most densely. This study explored the possibility that the male Harderian glands are a source of olfactory signals which promote proceptive behavior but suppress female agonistic behavior. Female gerbils in estrus were found to display significantly less than normal rates of proceptive behavior toward Harderianectomized males. The proceptive activity which was observed appeared to be slowed, but the typical pattern was retained. Female aggression, however, was not affected by their estrous condition or by the Harderian state of the males. Possibly Harderian letdown in male gerbils may inform females as to the reproductive competence of the males.     

Courtship feeding in Drosophila subobscura. II. Courtship feeding by males influences female mate choice

Male Drosophila subobscura provide females with a drop of regurgitated food during courtship. Males that were physically prevented from producing this drop (1) had reduced courtship success with starved, but not with fed, females; (2) positioned themselves in front of the female less frequently than males providing drops; and (3) had a smaller proportion of frontal displays which led to copulation. Males that produced small or watery drops also had reduced courtship success with starved, but not with fed, females. The starved females moved away from such males more frequently than from males providing large or yeasty drops. Evidence is presented in support of the hypothesis that the drops of food offered by males during courtship slow the moving female down, making it easier for the male to complete a frontal display, and to circle and attempt to mount before the female moves away after taking the food. The effectiveness of the drop in this role depends on its attractiveness to the female, which is influenced by the drop's size and content and by the female's nutritional status.