The effect of body coloration and group size on social partner preferences in female fighting fish (Betta splendens)

Females of the fighting fish Betta splendens have been shown to associate with other B. splendens females in a manner reminiscent of shoaling behavior. Since body coloration varies dramatically in this species, and since body coloration has been shown to affect shoalmate choice in other species of fish, we examined the influence of body coloration on association preferences in female B. splendens. In dichotomous choice tests, B. splendens females spent more time swimming near groups of females (regardless of coloration) than swimming near an empty chamber, and chose to swim near fish of similar coloration to their own when choosing between two distinctly colored groups of females. When examining the interplay between body coloration and group size, focal fish spent more time swimming near larger groups (N = 5) of similarly colored fish than swimming near an individual female of similar coloration. However, focal fish showed no preference when presented with an individual female of similar coloration and a larger group of females of dissimilar coloration. These results suggest that association choices in B. splendens females are strongly affected by both body coloration and by group size.     

光周期对雄性布氏田鼠种内个体气味辨别的影响

本文研究了成年雄性布氏田鼠对不同光照周期下(长光照: LD; 短光照: SD) 的陌生雌鼠和陌生雄鼠气味的行为表现。实验发现: 所有被试雄鼠对LD 雄鼠、雌鼠气味比SD 雄鼠、雌鼠气味有更多的社会探究行为。LD 雄鼠比SD 雄鼠对同性或异性陌生个体气味的嗅闻和挖掘行为要多, 而且差异显著。在探究LD 动情雌鼠气味源时, LD 雄鼠比SD 雄鼠表现出更多的嗅闻和挖掘行为; 在探究SD 动情雌鼠气味源时, LD 和SD 雄鼠的行为反应没有明显差别。所有被试雄鼠在LD 雄鼠气味源箱中的停留时间都显著多于在SD 雄鼠气味源箱中的停留时间。LD 雄鼠在LD 雌鼠气味源箱中的停留时间显著多于在SD 雌鼠气味源箱中的停留时间; 而SD 雄鼠在LD 和SD 雌鼠气味源箱中的进入频次和停留时间没有明显差别。同时, LD 雄鼠对LD 动情雌鼠的气味在嗅闻频次和嗅闻时间上多于非动情LD 雌鼠, 而对SD 动情和非动情雌鼠的气味没有表现出明显的偏好; SD 雄鼠对LD 和SD 动情雌鼠的嗅闻行为仅在频次上显著多于对非动情雌鼠的嗅闻, 而在时间上没有显著差别。结果表明: 布氏田鼠嗅觉通讯中的个体气味及其对气味进行辨别过程中的行为反应都随着光照周期的不同而发生变化。动物的个体气味带有季节性信息, 来源于长光照下的气味比短光照下的气味更具有性吸引; 短光照在一定程度上可能抑制了雌鼠的动情和雄鼠的性行为反应。其嗅觉通讯行为的适应意义可能在于: 减少秋季短日照环境下的繁殖活动, 提高雄性个体之间的社会容忍性以利于集群越冬。     

Social interactions among female rhesus macaques (Macaca mulatta) during the nonbreeding season: Responses to the introduction of males

When rhesus monkeys are observed in social groups during the breeding season, increases in interfemale aggression coincide with midcycle increases in sexual activity between males and females. However, some investigators have suggested that both aggressive and affiliative interactions between females are influenced by the presence or absence of males, irrespective of menstrual cycle stage. In the present study, social interactions among members of a captive group of rhesus females were measured during the non-breeding season in response to the introduction of rhesus males. Ovariectomized rhesus females (estrogen-treated or untreated) served as stimulus controls. Tests with males were characterized by significantly decreased interfemale proximity and grooming and significantly increased aggression from that seen in tests with stimulus females or in the absence of stimulus animals. Only interfemale proximity declined significantly during stimulus female tests, but results suggest that this may merely reflect a decline in this behavior that occurs across the course of the day. Estrogen treatment did not alter either the aggressive or affiliative behavior of stimulus females or group female response to stimulus females. The possibility is discussed that changes in interfemale interactions during tests with males reflect female interest in interacting with the male, particularly under social conditions that may limit such interactions.     

Who's Watching Me: Female Siamese Fighting Fish Alter Their Interactions in Response to an Audience

It is well established that interactions between conspecifics are often influenced by the presence of passive bystanders. Individuals have been found to alter their behavior in a variety of contexts, from foraging to aggression, based on the presence, sex, or identity of an audience. This audience effect may influence not only the nature of a signaling event but also the evolution of signal structure as signals may have to convey information across a distance. Additionally, audience individuals may use information obtained by watching in later encounters with these individuals, which may act as a selection pressure on communication. Communication networks in Siamese fighting fish, Betta splendens, are particularly well studied, with audience effects influencing both male–male interactions and male–female interactions. However, the effects of an audience on female–female interactions have not been examined in this species or any other. This study examined the interactions of pairs of females in three different audience conditions (male, female, and no audience). The results suggest that female–female interactions are affected by the presence of an audience as interactant‐directed gill flaring, the most commonly performed behavior, was performed more with an audience present. Additionally, the sex of the audience seemed to be influential, reflected by a difference in the frequency of interactant‐directed behaviors when a female vs. a male audience was present. This study is one of the first to demonstrate that females modify their behavior as a result of being watched and stresses the importance of examining audience effects in a variety of social contexts.     

Effects of Pair-Bond and Social Context on Male–Female Interactions in Captive Titi Monkeys (Callicebus moloch, Primates: Cebidae)

In monogamous species, an abiding relationship between a specific adult male and a specific adult female is a defining feature of the social system. The interactions between these individuals are influenced by many factors, including not only the history of their relationship (for example, development of a mutual bond), but also the immediate effects of the prevailing social context (for example, presence and sex of extra‐pair conspecifics). In this study we examined the effects of an existing bond and of social context on interactions between adult heterosexual pairs of the monogamous titi monkeys (Callicebus moloch). Twelve adult males and 12 adult females were tested with their cagemates and with an unfamiliar partner of the opposite sex in five social contexts: (1) mated male–female pair; (2) unfamiliar pair; (3) single female; (4) single male; and (5) empty stimulus cage. Results show that mated pairs were more affiliative than unfamiliar pairs and differentiated social contexts more sharply. Males were more responsive to context than females. Distance between mates was less and physical contact was more frequent in the presence of male–female pairs or a single male, than in the presence of a single female or an empty cage. These findings suggest that the presence and sex of strangers have a stronger influence on male–female interactions when the pair has an existing relationship.     

One-male harems and female social dynamics in Guinea baboons

Little is known about the mating system and social organization of Guinea baboons. This study investigated whether Guinea baboons have a harem-based mating system similar to that of hamadryas and gelada baboons and whether one-male mating units also correspond to social units. Ten adult females in a captive multi-male multi-female group of Guinea baboons were focally observed 2 h per week for 12 weeks, and all observed copulations within the group were recorded. Some males copulated with a single female while others had harems of 2-4 females. All females copulated with a single male except 1 female that switched harems early in the study. The focal females had higher rates of social interaction with their harem members, especially their harem male, than with individuals outside the harem. Females appeared to be subordinate to the harem male but little or no physical aggression or herding behavior from the male was observed. Variation in female social interactions within the harem was not accounted for by their sexual interactions with the male or their genetic relatedness with the females. Females, however, appeared to maintain social relationships with their female relatives in other harems. Taken together, the results of this study show that both mating and affiliative interactions in Guinea baboons are concentrated within one-male units and that the social dynamics within and between these units share some similarities as well as differences with those of hamadryas and gelada baboons.     

Microsatellite evidence for sperm storage and multiple paternity in the marine gastropod Crepidula coquimbensis

In gregarious species with copulation and internal fertilization, male-male competition and female cryptic choice may affect reproductive success of both sexes. We carried out a molecular analysis to study paternity and sperm use by females in the protandrous marine brooding gastropod Crepidula coquimbensis. In the field, a single female inhabits an empty hosting shell with up to six males. This gregarious behavior may promote intra-brood multiple paternity if females can store sperm from several consecutive copulations with the surrounding males. To study female sperm usage, the males sharing shelters with five different adult females were collected and preserved for paternity analysis. Females were transported alive to the laboratory and isolated for six months. After that, an additional male was offered to each of the five study females. Once the females had laid capsules, a total of 528 embryos from the five females were assigned paternity based on five microsatellite loci. Paternity analysis showed that every male sharing the empty hosting shell of a female as well as the additional male were assigned fatherhood of embryos laid by this specific female. Females can thus use sperms from multiple males including sperms stored for at least six months. In addition, in four out of the five offspring arrays, a similar contribution of each male to the brood was observed, a pattern associated with the close relationship between the number of fathers observed and the effective paternity index calculated. These results contrast with those of paternity analyses carried out in another species of the same genus, C. fornicata which is characterized by a stacking behavior in which the closest male to the female achieves the highest reproductive success. Male reproductive success may be largely influenced by the aggregation pattern and male mating opportunities in the Crepidula complex, a hypothesis to be examined further by studying other species exhibiting different grouping behavior.     

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.     

Audience Type and Receptivity Affect Male–Female Interactions in Siamese Fighting Fish

The study of communication in a network setting has gained increasing popularity in recent years. While audience effects on aggressive interactions have been studied extensively, male–female interactions have often been overlooked. In addition, little is known about how reproductive status affects the nature of audience effects. Siamese fighting fish, Betta splendens, are a popular subject for communication network studies, but male–female interactions have not been explored in this setting. In this study, pairs of male and female Betta were presented with a male, female or no audience to determine whether the presence of an audience alters the behaviour of the interactants. Within these three audience types, there were four reproductive status conditions with receptivity indicated by nest presence for males and reproductive barring for females. It was predicted that male–female interactions would be affected by the presence of an audience, especially when both interactants are receptive as has been found in male–male interactions in this species. The results suggest that presence of an audience and reproductive status act in combination to influence male–female interactions, but only in interactant‐directed behaviours. Not all behaviours were equally affected by these factors. For example, while tail beats to the other interact were greatest when a female audience was present and both the interactants were receptive, this was not true for gill flaring. This study is among the first investigations into audience effects on male–female interactions including the first in Betta and suggests that courtship as well as aggression should be explored in a network setting.     

Relation between the dominance rank of female rhesus monkeys and their access to males

Systematic observations were made on 12 measures of the sexual, aggressive, and social interactions of 24 male–female pairs of rhesus monkeys in six social groups, each consisting of one male and four ovariectomized females tested in a large room. Each female in a group was treated in turn first with estradiol alone and then with estradiol and progesterone in combination. When hormone-treated, the female was also observed during pair tests with the male in the same large observation room (four males, eight females, 240 group tests, 240 pair tests). The dominance ranks of females during group tests were determined post hoc by means of the dominance index [Zumpe & Michael, American Journal of Primatology 10:291–300, 1986]. In all six groups, the most dominant female virtually monopolized the male, and the subordinate females' interactions with the male, assessed during pair tests, were almost completely suppressed during group tests. This “dominant female effect” was a robust phenomenon that depended solely on female dominance rank. It was independent of the identity and hormonal status of females and of the social preferences of males as expressed in pair tests. These findings demonstrate the existence of female mate competition in an Old World primate.     

Interactions of social,spatial and hormonal factors on the behavior of rhesus monkeys (Macaca mulatta)

In a three year study, the interactions of social, spatial and hormonal variables on the behavior of rhesus monkeys were investigated. Observations were made on (1) an oppositelysexed pair alone in a cage; (2) the pair alone in a large room; and (3) the pair together with three ovariectomized females in the same large room. Each of four males was tested with the same group of four ovariectomized females (16 pairs, 480 1-hr tests). Each female served in turn as the focal female, namely, the hormone-treated one, and received first estradiol benzoate (EB) alone and then in combination with progesterone (P). The presence of three other females (the social variable) had a marked effect on all aspects of the behavior of the focal pair, and reduced the male's ejaculations as well as other sexual and affiliative interactions with the focal female. This was due to an inhibitory influence exerted by the dominant female on the interactions of subordinate females with the male. Comparing data from two groups of females showed that female dominance rank, and not male sexual preference, was the critical factor, and that mate competition between female rhesus monkeys was operating. Decreasing the space available for the interactions of the pair appeared to imitate a bonding effect in the male, increasing male grooming and tending to increase male sexual activity. In the group setting, treating the female with EB increased sexual activity, and additional progesterone decreased it in all three test settings. The effects of progesterone were most clearly apparent when the pair was tested alone in a large area.     

Sexual and social competition: broadening perspectives by defining female roles

Males figured more prominently than females in Darwin's view of sexual selection. He considered female choice of secondary importance to male-male competition as a mechanism to explain the evolution of male ornaments and armaments. Fisher later demonstrated the importance of female choice in driving male trait evolution, but his ideas were largely ignored for decades. As sexual selection came to embrace the notions of parent-offspring and sexual conflict, and experimental tests of female choice showed promise, females began to feature more prominently in the framework of sexual selection theory. Recent debate over this theory has centred around the role of females, not only over the question of choice, but also over female-female competition. Whereas some have called for expanding the sexual selection framework to encompass all forms of female-female competition, others have called for subsuming sexual selection within a broader framework of social selection, or replacing it altogether. Still others have argued for linking sexual selection more clearly to other evolutionary theories such as kin selection. Rather than simply debating terminology, we must take a broader view of the general processes that lead to trait evolution in both sexes by clearly defining the roles that females play in the process, and by focusing on intra- and inter-sexual interactions in males and females.     

Male and Female Mating Behavior is Dependent on Social Context in the Butterfly Bicyclus anynana

Reproduction is often more costly to females than it is to males, leading to the evolution of ornamented or competitive males and choosy females. Reproduction costs to females, however, can be reduced through nuptial gifts provided by males. These gifts, by increasing female survival or fecundity, can promote the evolution of mutual mate choice, ornamentation, or competition in both sexes, as well as plasticity in mating behavior dependent on social context. We tested for plasticity in male and female mating behavior in a species of butterfly, Bicyclus anynana, where male spermatophore gifts contribute to female survival and fecundity, and where mutual mate choice and ornamentation were previously established. We examined the effect of a sexual competitor on male–female interactions by observing and comparing the behavior of male–female pairs with that of triads containing either an extra male or an extra female. In the presence of a sexual competitor both males and females copulated less than when in male–female pairs, regardless of the direction of sex-ratio skew. Active males increased their own likelihood to copulate, while active females increased their likelihood of being courted. In addition, there was an effect of social context on relative rates of male and female courting and flying. These results suggest that both males and females change their mating behavior in response to social context in the butterfly Bicyclus anynana.     

Female contributions to the peaceful nature of bonobo society

Although chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) are closely related, females of the two species show surprisingly large differences in many behavioral aspects. While female chimpanzees tend to range alone or in small parties during non-estrous periods, female bonobos aggregate even more often than do males. Female chimpanzees do not have frequent social interactions with other females, whereas female bonobos maintain close social associations with one another. Although the ranging patterns of chimpanzee parties are generally led by males, female bonobos often take the initiative in ranging behavior. While female chimpanzees usually do not exhibit estrus during postpartum amenorrhea or pregnancy, female bonobos exhibit a prolonged pseudo-estrus during such non-conceptive periods. Studies of these two species have also shown great differences in agonistic behaviors performed by males. Male chimpanzees frequently fight with other males to compete for estrous females, but male bonobos seldom do so. While there are many records of infanticide by male chimpanzees, there is no confirmed record of such an event among bonobos. Several cases of within-group killing among adult male chimpanzees have been reported, but there is no such record for bonobos. While intergroup conflicts among chimpanzees sometimes involve killing members of the other group, intergroup conflicts among bonobos are considerably more moderate. In some cases, bonobos from two different groups may even range together for several days while engaging in various peaceful interactions. I will address two important questions that arise from these comparisons, exploring why females of such closely related species show such clear differences in behavior and whether or not the behavioral characteristics of female bonobos contribute to the peaceful nature of bonobo society.     

Testosterone increases singing and aggression but not male-typical sexual partner preference in early estrogen treated female zebra finches

Female zebra finches given estradiol benzoate (EB) as nestlings and testosterone propionate (TP) as adults show masculinized sexual partner preference, preferring females instead of males. This suggests an organizational effect of EB on sexual partner preference in a socially monogamous species that pairs for life. It is not known whether there is an activational hormone effect on sexual partner preference in this species, or whether adult testosterone treatment is necessary for masculinized preference to be expressed. In this experiment females were injected with EB daily for the first 2 weeks posthatching. As adults they were given TP filled or empty implants. Subjects were then given two-choice preference tests with male vs female stimuli, in which singing as well as proximity to the stimuli was recorded, followed by tests in a group aviary for social behavior and pairing preference. Females with TP implants sang more than females with empty implants and were more aggressive toward other females. They did not, however, differ from females with empty implants in any measure of sexual partner preference. Neither group showed a marked preference for males; instead both groups were equally interested in males and females. Thus adult testosterone treatment is not necessary for early estrogen treated females to show a shift in sexual partner preference in the male-typical direction.     

Mate choice copying in the sailfin molly, Poecilia latipinna, in the wild

Mate choice copying has been documented extensively in the laboratory with almost no supporting data available from studies in the wild. We investigated male and female mate choice copying in a wild population of the sailfin molly, a species that shows copying in the laboratory. We set up two upside-down plastic tanks in a river, with two jars of water on each tank. In male mate choice trials we placed a female in one jar and a male in the other on one tank and a female in one jar on the other tank, leaving the last jar empty. In female mate choice trials we presented a male and a female on one tank and a male and an empty jar on the other. Males preferred to associate with a female adjacent to a male rather than a lone female and females preferred to associate with a male adjacent to a female rather than a lone male. In two controls for shoaling behaviour we presented two males on one side of the set-up and one male on the other or two females versus one female. These controls showed that shoaling behaviour could not explain the male and female preference. Thus both sexes of the sailfin molly show mate choice copying in the wild, much as they do in laboratory studies. At least in this species, mate choice copying is not a laboratory artefact. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Shoaling in juvenile guppies: the effects of body size and shoal size

While factors affecting shoal mate choice have been examined extensively in adult guppies (Poecilia reticulata), few studies have focused on the shoaling behavior of juveniles. In this study, juvenile guppies were tested for their ability to shoal as well as their response to shoal mates of different body size and to shoals with different numbers of individuals. In dichotomous choice tests, 10-day-old guppies (mean body length=8.83 mm), 30-day-old guppies (13.17 mm) and 50-day-old guppies (18.6mm) were given the opportunity to swim near shoals of five fish or an empty chamber. In most cases, the juvenile fish demonstrated shoaling behavior, swimming near a group of fish rather than an empty chamber, regardless of the age of the stimulus shoal. When presented with two shoals, one of similar age and body size and one of dissimilar age and body size, only the 50-day-old guppies showed a significant preference for the age-matched shoal. Similarly, when choosing between a large shoal and a small shoal, only the 50-day-old guppies spent significantly more time near the larger shoal. Thus, while juveniles at each age shoaled, only 50-day-old fish demonstrated the shoal mate discrimination seen in adult fish.     

Sexual discrimination in the hermit crabPagurus geminus

Factors concerning sexual discrimination and related sensory receptors were investigated on the hermit crabPagurus geminus. In the breeding season, male crabs of this species were observed to hold receptive females but to reject males. By using this behavior, it was examined whether males on which some operation on the sensory receptors was performed could discriminate between presented male of female crabs. Test males could not discriminate sexes of the presented crabs only when antennules were removed bilaterally. On the other hand, intact test males reacted positively to an empty shell from which “female water” exuded after passing through the chamber containing females. Thus, the factor by which the hermit crab discriminates sexes was indicated to be a water-borne chemical and that the sensory receptor related to sex-discrimination appeared to exist on the antennules. Discussions are made with respect to predicted substances taking part in sexual discrimination of this hermit crab.     

Concordance in mate choice in female mound-building mice

Females must evaluate male quality to perform mate choice. Since females generally base their selection on different male features, individual females may differ in their choice. In this study, we show that concordance between females in mate choice decisions may arise without any experimental maximization of a particular attractive trait. Choice tests were performed in mound-building mice, Mus spicilegus, a monogamous species. Body odours of two male donors were presented to 12 female subjects individually. To determine female choice, the same pair of males was presented three times to a female. Four different pairs of male body odours were used. Male donors, not related to females, were selected at random in our polymorphic breeding stock. Using this two-way choice design, female mice displayed a clear choice and had a similar preference for particular males.