Male sexual harassment alters female social behaviour towards other females

Male harassment of females to gain mating opportunities is a consequence of an evolutionary conflict of interest between the sexes over reproduction and is common among sexually reproducing species. Male Trinidadian guppies Poecilia reticulata spend a large proportion of their time harassing females for copulations and their presence in female social groups has been shown to disrupt female-female social networks and the propensity for females to develop social recognition based on familiarity. In this study, we investigate the behavioural mechanisms that may lead to this disruption of female sociality. Using two experiments, we test the hypothesis that male presence will directly affect social behaviours expressed by females towards other females in the population. In experiment one, we tested for an effect of male presence on female shoaling behaviour and found that, in the presence of a free-swimming male guppy, females spent shorter amounts of time with other females than when in the presence of a free-swimming female guppy. In experiment two, we tested for an effect of male presence on the incidence of aggressive behaviour among female guppies. When males were present in a shoal, females exhibited increased levels of overall aggression towards other females compared with female only shoals. Our work provides direct evidence that the presence of sexually harassing males alters female-female social behaviour, an effect that we expect will be recurrent across taxonomic groups.     

Social preferences based on sexual attractiveness: a female strategy to reduce male sexual attention

Male sexual harassment of females is common across sexually reproducing species and can result in fitness costs to females. We hypothesized that females can reduce unwanted male attention by constructing a social niche where their female associates are more sexually attractive than themselves, thus influencing the decision-making of males to their advantage. We tested this hypothesis in the Trinidadian guppy (Poecilia reticulata), a species with high levels of male sexual harassment. First, we confirmed that non-receptive females were harassed less when they were paired with a more sexually attractive (receptive) female than with another non-receptive female. We then found that, indeed, females exploit this as a strategy to reduce sexual harassment; non-receptive females actively preferred to associate with receptive over non-receptive females. Importantly, when given access only to chemosensory cues, non-receptive females still showed this preference, suggesting that they use information from chemical cues to assess the sexual attractiveness of potential female partners. Receptive females in contrast showed no such preferences. Our results demonstrate that females can decrease male harassment by associating with females that are more sexually attractive than themselves and that they perform active partner choices based on this relative attractiveness. We propose that this strategy is likely to represent an important pathway by which females can construct social niches that influence the decision-making of others to their advantage; in this case, to reduce the sexual harassment they experience.     

Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition

Across sexually reproducing species, males and females are in conflict over the control of reproduction. At the heart of this conflict in a number of taxa is male harassment of females for mating opportunities and female strategies to avoid this harassment. One neglected consequence that may result from sexual harassment is the disruption of important social associations. Here, we experimentally manipulate the degree of sexual harassment that wild female guppies (Poecilia reticulata) experience by establishing replicated, semi-natural pools with different population sex ratios. We quantify the effects of sexual harassment on female social structure and the development of social recognition among females. When exposed to sexual harassment, we found that females had more disparate social networks with limited repeated interactions when compared to females that did not experience male harassment. Furthermore, females that did not experience harassment developed social recognition with familiar individuals over an 8-day period, whereas females that experienced harassment did not, an effect we suggest is due to disruption of association patterns. These results show that social network structure and social recognition can be affected by sexual harassment, an effect that will be relevant across taxonomic groups and that we predict will have fitness consequences for females.     

Predators, reproductive parasites, and the persistence of poor males on leks

Lekking males are thought to face strong directional selectionon secondary sexual traits. How variation in male traits canpersist under these conditions remains problematic (the lekparadox). Here, we present several game-theoretic models thatshow that avoidance of costly and mobile predators, sneakers,or brood parasites (enemies) leads to variation in female choice.This can result in maintenance of variation in male quality."Enemies" will congregate around higher quality males. Femalesmust then trade-off the benefits of mating with high-qualitymales against the increased risk of enemies. At equilibrium,the models predict a positive correlation between the qualityof a male and the proportions of both enemies and females visitinghim. In the first model, we use this framework to predict thelowest quality male on the lek that will receive any matings.In the second model, we examine the influence of this female-enemygame on the maintenance of variation in male quality. Low-qualitymales are likely to persist when enemies are costly to femalesor occur at high density, and when there is some spatial structureon the lek, so that neighboring males are typically of similarquality. If enemies are more costly to males than to females,high-quality males may benefit from receiving fewer female visits.In the third model, we consider the special case when enemiesare male reproductive parasites. These models illustrate theimportance of considering the simultaneous decisions of multipleplayers in mate choice games.     

Male harassment drives females to alter habitat use and leads to segregation of the sexes

Sexual conflict is ubiquitous across taxa. It often results in male harassment of females for mating opportunities that are costly for females, in some cases reducing reproductive success and increasing mortality. One strategy that females may employ to avoid sexual harassment is to segregate spatially from males. In fact, we do find sexual segregation in habitat use in species that have high levels of sexual conflict; however, the role of sexual harassment in driving such segregation remains poorly understood. Here, we demonstrate experimentally in a population of wild Trinidadian guppies Poecilia reticulata that male sexual harassment drives females into habitats that they otherwise do not prefer to occupy. In support of the social factors hypothesis for sexual segregation, which states that social factors such as harassment drive sexual segregation, this female behaviour leads to segregation of the sexes. In the presence of males, females actively select areas of high predation risk, but low male presence, and thus trade off increased predation risk against reduced sexual harassment.     

Phenotypic variation in male guppies from natural inland populations: an additional test of Haskins' sexual selection/predation hypothesis

Synopsis In Venezuela, guppies,Poecilia reticulata, are found in a diversity of aquatic ecosystems fringing the western slope of the eastern Andes. These inland drainages are associated with large between-site variation in both physical and biotic environments. Numbers of potential guppy predators vary greatly between low diversity piedmont streams and high diversity llanos swamps. We analyze geographic variation in several phenotypic traits associated with conspicuousness of male guppies as an independent test of Haskins' (Haskins et al. 1961) hypothesis of natural balance between sexual selection (driving male conspicuousness) and predation (driving crypsis). Four sites were compared: a diverse llanos swamp (high predation), two Andean piedmont streams (intermediate and low predation), and a spring pool located south of Venezuela's Paría Peninsula (no aquatic predators). Because tropical wet and dry seasons are associated with varying degrees of environmental change at these sites, within-site seasonal samples provided an additional test of the role of selective predation on Male phenotypes. Phenotypic traits associated with both size and coloration supported the sexual selection/predation hypothesis. male guppies from the spring were generally more colorful than male guppies from inland drainages. Males from the diverse swamp site were smaller and less colorful than guppies from other sites. Males from the swamp population were smallest and least colorful during the early dry season, when diurnal aquatic piscivores were more diverse and abundant. Dominant spot pigments at each site were generally those that provided greatest contrast with the predominant environmental background color, lending further evidence that coloration patterns in male guppies enhance conspicuousness.     

Experimentally induced variation in neuroendocrine processes affects male reproductive behaviour,sperm characteristics and social interactions

While extensive research has focused on how social interactions evolve, the fitness consequences of the neuroendocrine mechanisms underlying these interactions have rarely been documented, especially in the wild. Here, we measure how the neuroendocrine mechanisms underlying male behaviour affect mating success and sperm competition in the ocellated wrasse (Symphodus ocellatus). In this species, males exhibit three alternative reproductive types. “Nesting males” provide parental care, defend territories and form cooperative associations with unrelated “satellites,” who cheat by sneaking fertilizations but help by reducing sperm competition from “sneakers” who do not cooperate or provide care. To measure the fitness consequences of the mechanisms underlying these social interactions, we used “phenotypic engineering” that involved administering an androgen receptor antagonist (flutamide) to wild, free‐living fish. Nesting males treated with flutamide shifted their aggression from sneakers to satellite males and experienced decreased submissiveness by sneaker males (which correlated with decreased nesting male mating success). The preoptic area (POA), a region controlling male reproductive behaviours, exhibited dramatic down‐regulation of androgen receptor (AR) and vasotocin 1a receptor (V1aR) mRNA following experimental manipulation of androgen signalling. We did not find a direct effect of the manipulation on male mating success, paternity or larval production. However, variation in neuroendocrine mechanisms generated by the experimental manipulation was significantly correlated with changes in behaviour and mating success: V1aR expression was negatively correlated with satellite‐directed aggression, and expression of its ligand arginine vasotocin (AVT) was positively correlated with courtship and mating success, thus revealing the potential for sexual selection on these mechanisms.     

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.     

接种丛枝菌根真菌对柑橘生长与次生代谢的影响

为了解丛枝菌根(AM)真菌对根、根系分泌物中次生代谢物和植物生长的影响,以AM真菌(Glomusepigaeum)接种柑橘(Citrusreticulata),对柑橘的酚类物质、抗氧化酶和生长情况进行研究。结果表明,温室盆栽6个月后,接种AM真菌显著提高柑橘酚酸类物质的积累,但是酚酸组分在根和根分泌物中存在差异,根中原儿茶酸和丁香酸含量为29.98和18.32μg/g,分别是未接种的4.58和2.26倍。而根系分泌物中香豆酸、苯甲酸和根皮苷含量为0.36、6.04和12.32μg/g,分别是未接种的1.71、1.94和1.25倍,而香草醛仅在未接种柑橘根中检测到。接种AM真菌的柑橘苯丙氨酸解氨酶、多酚氧化酶和过氧化物酶活性为38.36、0.51和28.62 U/(g·min),分别是未接种AM真菌的1.99、2.83和3.10倍。同时菌根定殖也显著提高柑橘的株高、茎粗和叶片数。因此,AM真菌定殖能促进植物生长,改变柑橘次生代谢产物的积累。     

端黑萤和边褐端黑萤两性异形和雌性个体生育力

检测了端黑萤(Abscondita chinensis)和边褐端黑萤(Abscondita terminalis)繁殖期个体大小和形态特征的两性异形以及雌性繁殖输出。两因素方差分析表明,端黑萤的全长显著小于边褐端黑萤的全长,雌性个体显著大于雄性个体。以全长为协变量的两因素协方差分析显示,端黑萤的前胸背板宽、鞘翅长、复眼宽、胸长、腹宽、发光器面积及体重均显著小于边褐端黑萤,而腹长显著大于边褐端黑萤,触角长物种间差异不显著;雌性的前胸背板宽、腹长、腹宽及体重均显著大于雄性,而鞘翅长、复眼宽、胸长、触角长、发光器面积均显著小于雄性;物种和性别的交互作用对前胸背板宽、复眼宽、腹长、触角长及发光器面积影响显著,对其余的鞘翅长、胸长、腹宽及体重影响不显著。端黑萤和边褐端黑萤的两性异形指数分别为0.144和0.091。9个形态特征变量的主成分分析(Eigenvalue ≥ 1)发现,前2个主成分共解释56.8%的变异;复眼宽和发光器面积在第一主成分有较高的负负载系数,腹长有较高的正负载系数(解释31.3%变异);前胸背板宽和鞘翅长在第二主成分有较高的负负载系数(解释25.5%变异)。端黑萤和边褐端黑萤的平均窝卵数分别为28.9粒和18粒。线性回归显示,端黑萤和边褐端黑萤的窝卵数均与母体的全长及体重呈显著的正相关。单因素方差分析表明,特定全长的端黑萤的窝卵数显著大于边褐端黑萤。端黑萤和边褐端黑萤均属于雌性偏大的两性异形,是生育力选择、能量分配和运动综合影响的结果。雌萤腹腔容量等关键局部特征的增大是对生育力选择的适应,且生育力大小与两性异形差异程度成正相关。