Cave molly females (Poecilia mexicana, Poeciliidae, Teleostei) like well-fed males

We examined the preference of Atlantic molly females (Poecilia mexicana) to associate with a well-fed or a starved male in simultaneous choice tests. Females from three different populations were tested in three treatments: (1) the females could choose on the basis of multiple cues from the males (visual plus non-visual); (2) only non-visual cues could be perceived in darkness, (3) only visual cues were presented. The three tested populations differ clearly in their ecology: one population occurs in a typical river habitat, the second one in a milky sulfur creek outside a cave, and the third population occurs in a cave habitat (cave molly). In the river-dwelling population, females never showed a preference. In the population from the sulfur creek, females preferred to associate with the well-nourished male when visual cues from the males were available. Only cave molly females exhibited a strong preference for well-nourished males in all treatments. A morphological comparison demonstrated that wild-caught males from river habitats are typically in a good nutritional state. In the sulfur creek, males showed signs of starvation. Cave molly males were in an even worse nutritional state. In the cave population, saturated males probably indicate high fitness, thereby driving the evolution of the preference for good male nutritional state.Communicated by K. Lindström     

Sexual selection in darkness? Female mating preferences in surface- and cave-dwelling Atlantic mollies,<Emphasis Type="Italic"> Poecilia mexicana</Emphasis> (Poeciliidae,Teleostei)

In many species, females prefer large males as mating partners. Male size can easily be determined visually. We examined how such mate choice evolves when individuals are deprived of sensory information previously used to determine male quality. Using a cave-dwelling population of Atlantic mollies, Poecilia mexicana, as a model, we asked whether the female mating preference for large male body size still occurs in this cave fish, or whether it is lacking due to the absence of visual communication in the cave environment. In simultaneous choice tests we compared the time females spent with a large or a small male. In the river-dwelling and cave-dwelling fish, as well as the population originating from the cave entrance, we found a significant preference for large males in light. Hence, the ancestral mating preference of surface-dwelling mollies has been maintained during the evolution of the cave form. However, only the cave population exhibited a mating preference in darkness. Determination of male body size appears to have been taken over by a non-visual sensory system, probably the lateral line.Communicated by K. Lindström     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

When is a male not a male? Sex recognition and choice in two sex-changing species

For dioecious species, choosing a mate of the same sex can have reproductive costs. For sex-changing animals, however, a lack of sex recognition may not carry a reproductive cost, as pairs that were initially same-sex can become opposite-sex pairs as one partner changes sex. The strength of sex discrimination in sex changers, then, should depend on the duration of mating associations and whether the time of sex change is influenced by social situation (“flexible” sex change). We studied two species of marine snails that change sex from male to female, one with flexible sex change and long-term or permanent mating associations (Crepidula fornicata) and one with short-term pairings and relatively fixed time of sex change (Crepidula convexa), to determine whether either species exhibits sex recognition and whether members of C. convexa show stronger sex discrimination. In laboratory experiments, small males, the choosing animals, were placed with either a male or a female conspecific (no-choice experiments) or given a choice of a male or female (choice experiments). We controlled for shell length in all experiments, as relative size may influence sex change or choice. Males of both species paired more often with females than males, but, as predicted, males of C. convexa showed stronger discrimination: When given a choice, no C. convexa male paired with another male. In contrast, some C. fornicata males always chose other males even when given the choice of a female. These results suggest that sex recognition can be adaptive even for sex changers but demonstrate that the level of sex recognition will depend on other aspects of reproductive behavior.     

Demonstration of sex pheromones in the predaceous diving beetle Rhantus suturalis (MacLeay 1825) (Dytiscidae)

Sex pheromones are used by a wide variety of species in terrestrial ecosystems. Much less is known about these pheromones in aquatic systems e.g., for diving beetles. To test the use of pheromones and visual signals for mate finding by the diving beetle Rhantus suturalis (MacLeay 1825), behavioral experiments were performed using three different types of vessels containing conspecific. In experiments with non-permeable glass flasks, which did not allow the diffusion of chemical substances, males and females did not stimulate any reaction by conspecific of either sex. In experiments with opaque but water permeable vessels, and in experiments with vessels made of finely woven steel, which allowed the perception of chemical as well as optical cues, male diving beetles were significantly attracted to females. Females were attracted to other females only in the last experiment, when simultaneously perceiving chemical and optical cues. These results indicate that R. suturalis females release sex pheromones to attract mating partners, which is the first experimental demonstration of pheromones in the Dytiscidae.     

Mate preferences in <Emphasis Type="Italic">Drosophila</Emphasis> infected with <Emphasis Type="Italic">Wolbachia</Emphasis>?

The maternally inherited bacterium Wolbachia pipientis generates strong reproductive incompatibilities between uninfected females and infected males (cytoplasmic incompatibility), significantly reducing both female and male reproductive success. Such fitness costs are thought to place selective pressure on hosts to evolve pre-copulatory preferences for mating with compatible mates, thereby enabling them to avoid the reproductive incompatibilities associated with Wolbachia. Therefore, uninfected females are predicted to prefer mating with uninfected males, whereas infected males are predicted to prefer mating with infected females. Despite these predictions, previous investigations of pre-copulatory mate preferences in Wolbachia-manipulated Drosophila have not found evidence of female preference for uninfected or compatible males. However, none of these studies utilised a design where focal individuals are provided with a simple choice in a relatively non-competitive situation. We examined both female and male pre-copulatory mate preference based on mate infection status in Drosophila simulans and D. melanogaster using simple choice assays involving between 30–50 replicates per treatment. Although we found no evidence of female pre-copulatory mate preferences in either species, male D. simulans exhibited some preference for mating with females of the same infection status. However, this preference was not evident when we repeated the experiment to confirm this finding. Consequently, we conclude that neither male nor female D. melanogaster and D. simulans exhibit significant Wolbachia-associated pre-copulatory mate preferences.     

Audience effect alters male but not female mating preferences

Males often face strong mating competition by neighboring males in their social environment. A recent study by Plath et al. (Anim Behav 75:21–29, 2008a) has demonstrated that the visual presence of a male competitor (i.e., an audience male) affects the expression of male mating preferences in a poeciliid fish (Poecilia mexicana) with a weaker expression of mating preferences when an audience male observed the focal male. This may be a tactic to reduce sperm competition, since surrounding males likely share intrinsic preferences for female traits or copy mate choice decisions. Here, we examined the hypothesis that a same-sex audience would affect female mate preferences less than male mating preferences. Our hypothesis was based on the assumptions that (1) competition for mates in a fashion that would be comparable in strength to sperm competition or overt male–male aggression is absent among Poecilia females, and (2) P. mexicana females typically form female-biased shoals, such that almost any female mate choice in nature occurs in front of a female audience. Poecilia females (P. mexicana, surface and cave form, and the closely related gynogenetic Poecilia formosa) were given a choice between a large and a small male, and the tests were repeated while a conspecific, a heterospecific, or no audience female (control) was presented. Females spent more time in the neutral zone and, thus, less time near the males during the second part of a trial when an audience was presented, but—consistent with predictions—females showed only slightly weaker expression of mate preferences during the second part of the tests. This decline was not specific to the treatment involving an audience and was significantly weaker than the effect seen in the male sex.     

Leg ornamentation and the efficacy of courtship display in four species of wolf spider (Araneae: Lycosidae)

This study used both correlative and experimental video playback methods to test the hypothesis that the secondary sexual traits of male wolf spiders act to increase the efficacy of visual courtship displays. Direct observations of courtship of several lycosid genera and a review of the literature revealed a significant association between ornamentation and visual courtship displays. This suggests that the ornamentation may be playing the role of amplifier for a visual display. To test this hypothesis, male courtship behaviors of four Schizocosa species were experimentally manipulated using video-imaging techniques. Females of species with non-visually displaying, non-ornamented males (Schizocosa duplex and S. uetzi) did not increase in frequency of receptivity when tufts were added to conspecific males. In a species with a visual display and foreleg pigmentation (S. stridulans), the addition of foreleg tufts increased female receptivity. In a tufted species (S. crassipes), females tended to decrease their receptivity when male ornamentation was completely removed. In visually displaying species, ornamentation acts to increase female receptivity, supporting its role as an amplifier of a visual display. Received: 29 December 1997 / Received in revised form: 23 October 1999 / Accepted: 13 December 1999     

The avoidance response of fathead minnows to chemical alarm cues: understanding the effects of donor gender and breeding condition

Summary. All animals are vulnerable to predation at some point in their lives and consequently prey organisms often develop effective risk assessment systems. For many aquatic species predation risk assessment occurs through the use of olfactory cues, including predator odours and alarm cues from damaged or disturbed conspecifics. When aquatic species encounter conspecific alarm cues they may respond, or not, based on specific information including cue concentration, health and size of the conspecific donor and potentially the gender and breeding condition of the donor. Previous laboratory studies have demonstrated that fathead minnows (Pimephales promelas) fail to respond to the skin extracts of breeding male minnows. The purpose of the current study was to verify these early laboratory findings in the field as well as to further investigate the effect of female reproductive state and donor gender on the response of minnows to damage-release alarm cues. Our results indicate that male breeding condition has a significant effect on how minnows will respond to conspecific cues. Minnows showed avoidance of cues of female minnows and male minnows not in breeding condition, in comparison to cues of breeding male minnows and cues of male and female swordtails. Neither the gender of non-breeding minnows nor the reproductive state of female minnows influenced the avoidance of minnows to alarm cues.     

Reinforcement of male mate preferences in sympatric killifish species Lucania goodei and Lucania parva

Reinforcement occurs when reduced hybrid fitness leads to the evolution of a stronger prezygotic isolation. Populations sympatric with closely related species, where hybridization occurs, are predicted to have stronger mate preferences than allopatric populations. The reinforcement of male mate preference is thought to be rarer than the reinforcement of female preference, but this inference may be biased by the lack of studies on male preference. We tested male mate preferences from sympatric and allopatric populations of two closely related species of killifish: Lucania goodei and Lucania parva. We found that sympatric males had greater preferences for conspecific females than allopatric males. Furthermore, conspecific preferences in allopatric populations were weakest when these populations were geographically distant (>50?km) from those of heterospecifics. Our data suggest that reinforcement has contributed to male conspecific preference and speciation in Lucania.     

Female mate choice determines reproductive isolation between sympatric butterflies

Animal courtship rituals are important for species recognition, and a variety of cues might be utilized to recognize conspecific mates. In this paper, we investigate different species-recognition mechanisms between two sympatric butterfly sister species: the wood white (Leptidea sinapis) and Real’s wood white (Leptidea reali). We show that males of both species frequently court heterospecific females both under laboratory and field conditions. The long-lasting elaborate courtships impose energetic costs, since the second courtship of males that were introduced to two subsequent conspecific females lasted on average only one fourth as long as the first courtship. In this paper, we demonstrate that premating reproductive isolation is dependent on female unwillingness to accept heterospecific mates. We studied female and male courtship behavior, chemical signaling, and the morphology of the sexually dimorphic antennae, one of the few male traits visible for females during courtship. We found no differences in ultraviolet (UV) reflectance and only small differences in longer wavelengths and brightness, significant between-species differences, but strongly overlapping distributions of male L. sinapis and L. reali antennal morphology and chemical signals and minor differences in courtship behavior. The lack of clear-cut between-species differences further explains the lack of male species recognition, and the overall similarity might have caused the long-lasting elaborate courtships, if females need prolonged male courtships to distinguish between con- and heterospecific suitors.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Nestmate recognition in three species of stenogastrine wasps (Hymenoptera,Vespidae)

The capacity to recognise a conspecific intruder was investigated in Parischnogaster jacobsoni, Liostenogaster flavolineata and L. vechti, three species of primitively social wasps of the subfamily Stenogastrinae. Results of behavioural experiments carried out in the field showed that females of all three species react pacifically if presented with female nestmates, but aggressively reject an intruder from a conspecific colony. As L. flavolineata and L. vechti both build large clusters of nests, often very close to each other, the recognition capacity among females from different nests, but in the same conspecific cluster, was also investigated. Females of both species were more aggressive towards females from a different colony in the same cluster than towards their female nestmates. Additional experiments on L. flavolineata showed that there was no difference in reaction towards females from colonies nearer or further from the tested colony but within the same cluster, nor towards females from a different cluster. The capacity to recognise an alien conspecific nest containing immature brood was investigated in P. jacobsoni. Adult females of this species, invited to land on an alien nest which had experimentally been exchanged for their own, accepted the new nest and partially destroyed the immature brood. The behaviour of the females when they land on an alien nest, however, suggests that they do recognise the nest as foreign. Acceptance of foreign nests coupled with low immature brood destruction is probably due to the high energetic costs of egg-deposition and larval rearing in stenogastrine wasps. These results suggest that nestmate recognition in these wasps is very efficient, even though they belong to the most primitive subfamily of social wasps. Received: 16 April 1996/Accepted after revision: 9 August 1996     

To beg or to freeze: multimodal sensory integration directs behavior in a tadpole

Effective coordination of behaviors such as foraging and avoiding predators requires an assessment of cues provided by other organisms. Integrating cues from multiple sensory modalities may enhance the assessment. We studied cue integration by tadpoles of Oophaga pumilio, which live in small arboreal water pools. In this species, mothers periodically visit their tadpoles and feed them with unfertilized eggs. When mothers visit, tadpoles beg conspicuously by vibrating until fed. However, animals other than mother frogs including potential predators may visit water pools. Thus, when a visitor appears, tadpoles must use visitor cues to decide whether to beg or to remain inactive to avoid predation. To elucidate the cues that prompt these behaviors, we videotaped behavior of O. pumilio tadpoles in response to isolated and multimodal cues. Tadpoles swam more when exposed to visual or visual and chemical cues of adult O. pumilio but only exhibited begging when exposed to visual, chemical, and tactile cues together. Visual, chemical, and tactile cues from either male or female adult O. pumilio stimulated swimming and begging, but the same cues from similarly sized heterospecific frogs did not. Lastly, tadpoles exposed to a potential predator did not beg and swam less than tadpoles with no stimulus. Together, these findings suggest that O. pumilio tadpoles use multimodal cues to modulate swimming behavior accordingly in the presence of egg provisioners, predators, and other visitors and that tadpole begging is induced by multimodal cues of conspecific frogs such that tactile and perhaps chemical cues supplement visual cues.     

Association patterns of sailfin mollies (Poecilia latipinna): alternative hypotheses

Individuals may associate with each other due to a variety of selective forces, such as intra- and intersexual selection, and conspecific recognition. Previous studies have concluded that mate choice governs association behavior in polygynous species of fish. I examined whether mate choice underlies the preference for larger individuals by examining preference for association (time spent in proximity to a fish) not only between opposite-sex individuals but also between same-sex individuals of the live-bearing sailfin molly (Poecilia latipinna). Males and females from three size classes were tested with a large and a small object fish of the same and opposite sex. Females preferred to associate with larger over smaller males. Males also preferred to associate with larger over smaller females, as expected. The same female and male test fish also preferred to associate with larger over smaller fish of the same sex. Moreover, females demonstrated no significant difference in their strength of preference (large–small) when offered males or females. The same held true for males. When males and females were subsequently tested with one large male and one large female, females tended to prefer large males while males showed no significant preference for association based on sex. In another experiment, females were tested with a large female and a small male, and significantly preferred the former. These findings suggest that association patterns may have arisen under a variety of conditions, such as predation pressures, shoaling behavior, and associative preference behavior. The assumption that association behavior is a uniformly sufficient predictor of mate choice in fish needs to be re-examined for P. latipinna and other species. Received: 6 November 1998 / Received in revised form: 12 May 1999 / Accepted: 12 May 1999     

Pheromones inBlattella germanica andBlaberus craniifer (Blaberoidea): Glandular source,morphology and analyses of pheromonally released behaviours

Females ofBlattella germanica andBlaberus craniifer produce a volatile sex pheromone attractive at a distance for conspecific males. During the emission of the pheromone, females ofB. craniifer adopt a typical calling posture; we never observe such a stance inB. germanica females. For both species, the glandular structures responsible for pheromonal production are located on the female pygidium. InB. germanica, the thickness of the glandular epithelium is clearly correlated with the attractiveness of females, being maximum for 13 day old females. The functions of the pygidial glands are discussed in both species.     

Differential sperm expenditure by male sailfin mollies, Poecilia latipinna, in a unisexual–bisexual species complex and the influence of spermiation during mating

Selection should favor strategies that reduce costs associated with spermatogenesis. This is especially true when males are sympatric with closely related species, and must avoid heterospecific matings, as in the unisexual–bisexual species complex of mollies. Male sailfin mollies, Poecilia latipinna, are sexually parasitized by Amazon mollies (P. formosa), and produce more sperm in the presence of female sailfin mollies than in the presence of Amazon mollies. We tested the hypothesis that male sailfin mollies differentially expend sperm when mating with either conspecific or heterospecific females. We measured sperm expenditure by determining the amount of sperm males have remaining after mating. Male sailfin mollies had more sperm available after mating with female sailfin mollies than after mating with Amazon mollies. While this result could indicate higher sperm expenditure to Amazon mollies, males mating with female sailfin mollies had more sperm available after mating than their baseline sperm reserves. Spermiation, the last stage of spermatogenesis, could be triggered by physical contact with females, and could increase sperm availability during mating. We examined the relationship between sperm availability and the amount of time that males mated with females. We found that sperm availability increased as mating trial time increased with female sailfin mollies, but not with Amazon mollies. Spermiation in the presence of conspecific female stimuli could reduce physiological costs associated with spermatogenesis while increasing the amount and quality of sperm available for sperm competition. We suggest that future studies examining sperm priming and expenditure should consider the potential for spermiation.     

The lexicon of love: what cues cause size-assortative courtship by male garter snakes?

Cues that females use to select potential mates have attracted substantial research effort, but the criteria for male mate choice remain very poorly known. Red-sided garter snakes ( Thamnophis sirtalis parietalis) court and mate in large aggregations around overwintering dens in southern Manitoba, Canada. Both courtship and mating are size-assortative: small male snakes court small as well as large females, whereas larger males court only large females. This system provides a unique opportunity to assess the cues that males use in selecting mates, and in particular the mechanisms that generate a size-related shift in mate preference. Experiments in which we manipulated body sizes and scents showed that both vision and scent (sex pheromones) were important. Large males directed intense courtship only when the stimulus provided both visual and chemical (skin lipid) evidence of large body size. Small males were much less discriminating in both respects. Thus, size-assortative mating in this system is generated not by larger males excluding their smaller rivals from the largest females (as has been reported in other reptile species), but by a size-related shift in the visual and pheromonal cues that elicit courtship. Males of some species may thus show complex patterns of mate choice, with the cues that stimulate courtship differing even among males within a single population based on traits such as age or body size.     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication