Sex recognition in surface- and cave-dwelling Atlantic molly females (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae,Teleostei): influence of visual and non-visual cues

Cave fishes need to rely on non-visual senses, such as the sense of smell or the lateral line to communicate in darkness. In the present study, we investigated sex identification by females of a cave-dwelling livebearing fish, Poecilia mexicana (cave molly), as well as its surface-dwelling relatives. Unlike many other cave fishes, cave mollies still possess functional eyes. Three different modes of presentation of the stimulus fish (a male and an equally sized female) were used: (i) the stimulus fish were presented behind wire-mesh in light, allowing the focal female to perceive multiple cues, (ii) the experiment was carried out under infrared conditions, such that only non-visual cues could be perceived and (iii) the stimulus fish were presented in light behind transparent Plexiglas, allowing for the use of visual cues only. Females of all populations examined preferred to associate with the stimulus female in at least one of the treatments, but only when visible light was provided, suggesting that far-range sex recognition is limited or even absent in the cave molly under naturally dark conditions.     

Shoaling behaviour in a surface-dwelling and a cave-dwelling population of a barb <Emphasis Type="Italic">Garra barreimiae</Emphasis> (Cyprinidae,Teleostei)

We studied shoaling behaviour in a species of fish (Garra barreimiae) from Oman. We compared two populations (a surface-dwelling and a cave-dwelling population) with different theoretical costs and benefits of shoaling. We measured the tendency to associate with a shoal of conspecifics. The stimulus shoal was confined to (1) clear Plexiglas cylinders in light, (2) wire-mesh cylinders in light, or (3) wire-mesh cylinders in darkness. The surface form exhibited a strong preference for the stimulus shoal during the experiments in light, but also in darkness, when only non-visual cues from the shoal could be perceived. The cave form did not show a preference when solely visual cues were available (Plexiglas cylinder). When non-visual cues from the shoal could be perceived (wire-mesh), the cave form did show a preference to associate with the shoal, but the shoaling tendency was considerably weaker than in the surface form. The shoaling tendency has probably been genetically reduced in the cave form.     

Nonvisual mate choice in the Pyrenean mountain newt (<Emphasis Type="Italic">Euproctus asper</Emphasis>): females prefer small males

In many species, females prefer large males as mates. In this paper, we examined whether female Pyrenean mountain newts (Euproctus asper) would show a preference for large males in a simultaneous choice situation. When we tested for a visual preference relative to male body size using visual cues, females did not prefer large males. When chemical cues were presented, females did show a preference, but contrary to our prediction, they preferred the smaller of two males. E. asper males exhibit coercive mating behavior (amplexus), keeping females from feeding for several hours or even days, which may explain why females avoid large, physically stronger, and thus, more coercive males. The predominantly nocturnal lifestyle may explain why visual cues are less important for mate discrimination in this species.     

Diverging Cave- and River-Dwelling Newts Exert the Same Mate Preference in their Native Light Conditions

When colonizing a new habitat, populations must adapt their sexual behaviour to new ecological constraints. Because caves display drastically different conditions from surface habitats and cave animals are deprived from visual information, hypogean populations are expected to have modified their mate preference and signalling behaviour after cave colonization. Here, we experimentally examined the female preference and the sexual behaviour of brook newts Calotriton asper from different cave and river populations, either in light or in darkness. Our results suggest that females prefer large individuals in both hypogean and epigean populations, but that this preference is only expressed in the light conditions of their native habitat. Hence, some mate choice criteria would be maintained across genetically divergent populations and throughout dissimilar habitats. However, this sexual behaviour is likely to be expressed via a different sensory pathway in the different habitats, suggesting that a sensory shift has occurred in cave populations, enabling animals to communicate through a non-visual channel.     

Males Prefer Small Females in a Dichotomous Choice Test in the Poeciliid Fish Heterandria formosa

Male choice is expected to evolve when females differ in quality, even if male investment in each mating is low. The family Poeciliidae is an example of fishes in which males show little parental investment as they only provide sperm. Up until now, a preference for large females has been found in all species studied. Here we show that unexpectedly, males of the least killifish (Heterandria formosa) prefer to interact with small instead of large females in a dichotomous male choice test, even though large females are more fecund. During a free‐swimming choice experiment, males did not discriminate between females based on their size. We suggest that this unique preference for small females, or the lack of preference for large females, results from strong first male sperm precedence in this species. Smaller females are younger and therefore more likely to be virgin, which probably makes them more profitable mates for males. When presented with a virgin and a mated female of similar size, males showed no preference for either type. This suggests that males do not use pheromone cues to assess female mating status but that they are likely to use female size as a proxy for it.     

The influence of conspecific chemical cues on walking behavior of <Emphasis Type="Italic">Ixodes ricinus</Emphasis> males

The influence of female and male cues on the male searching behavior of the sheep tick, Ixodes ricinus (Acari: Ixodidae), was studied under constant laboratory conditions by means of a computerized video tracking system. Males were released into an open glass arena in which one half had been exposed for 1 h to 5 freely walking unfed females or five males. The obtained track data were analyzed to reveal if male walking pattern was random or affected by cues of conspecifics left on the arena surface. The results showed that males spent significantly more time and walked a significantly longer distance in the area with female cues compared to the control area. Moreover, the walking path of males was more tortuous on the cues half than the control half. In contrast, walking behavior of males was not affected by male cues in any observed parameter. These results provide the first evidence that unfed females leave sex-specific semiochemicals on a substrate during their walk and that these cues have an effect on male searching behavior. Since this effect was not observed when female cues were 24-h old, the semiochemicals seem to either evaporate or are subject to degradation. The biological significance of these findings for mate-searching in I. ricinus males is discussed.     

Friend or foe: behavioral responses to conspecifics in the northern scorpion, <Emphasis Type="Italic">Paruroctonus boreus</Emphasis> (Scorpionida: Vaejovidae)

We investigated the behavioral response of male Paruroctonus boreus to male and female pheromones of conspecifics. Adult male scorpions actively search for females during the mating season, but lack visual cues to direct their movement. Chemical cues (pheromones) are thought to be responsible for directing movement. In a series of laboratory experiments, in Y-mazes, males were exposed to areas that had contained males and females from the same population and areas that had contained conspecifics from different populations. Males spent significantly more time in areas that had contained females from the same population but did not show a preference for areas exposed to females from different populations. Males also significantly avoided areas exposed to males from the same population. Our results suggest that P. boreus has a sex-specific response to pheromones and that this response may be population-specific.     

Sexual harassment in a live-bearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>): influence of population-specific male mating behaviour

We investigated male sexual behaviour and the cost of sexual harassment, as measured by the reduction of female feeding time in the presence of a male, in a cave-dwelling population of Poecilia mexicana, in which sexual harassment does not occur naturally. We asked whether the lack of sexual harassment in this population is due to low sexual activity of the males, or low feeding motivation of the females. We experimentally increased the sexual activity of males or the females feeding motivation, or we used a combination of both treatments. Female feeding time was not lower in the presence of a male than in the presence of a female after sexual deprivation of the males or food deprivation of the females. Only in the combined experiment was female feeding time lower in the presence of a large male than in the presence of a small male, indicating a weak effect of sexual harassment by large males. Virgin females did not suffer a cost of sexual harassment, indicating that sexual experience does not cause the lack of sexual harassment in cave mollies. Males from a surface population, where sexual harassment occurs, significantly reduced the feeding time of cave-dwelling females even though these males exhibited surprisingly little sexual behaviour. The sexual activity of cave mollies did not correlate with male body size in any experiment, indicating that even after sexual deprivation, small cave molly males do not switch to the alternative mating behaviour known in surface-dwelling P. mexicana, where sexual activity is correlated negatively with male body size.     

Quantitative genetic models of sexual selection by male choice

There are many examples of male mate choice for female traits that tend to be associated with high fertility. I develop quantitative genetic models of a female trait and a male preference to show when such a male preference can evolve. I find that a disagreement between the fertility maximum and the viability maximum of the female trait is necessary for directional male preference (preference for extreme female trait values) to evolve. Moreover, when there is a shortage of available male partners or variance in male nongenetic quality, strong male preference can evolve. Furthermore, I also show that males evolve to exhibit a stronger preference for females that are more feminine (less resemblance to males) than the average female when there is a sexual dimorphism caused by fertility selection which acts only on females.     

Males,but not females,contribute to sexual isolation between two sympatric species of <Emphasis Type="Italic">Gambusia</Emphasis>

Reproductive isolation restricts genetic exchange between species. Various pre- and post-mating barriers, such as behavior, physiology and gametic incompatibility, have been shown to evolve in sympatry. In certain scenarios, isolation can be asymmetrical, where species differentially prefer conspecifics. We examined sexual isolation via conspecific mate preference between Gambusia affinis and G. geiseri in both sexes. To investigate male contribution to sexual isolation, we compared the number of mating attempts (gonopodial thrusts) directed at either a conspecific or a heterospecific female, in both species. We also examined sperm priming and expenditure in males in the presence of conspecific or heterospecific females. We then measured female preference for either a conspecific or heterospecific male, in both species. We found that males of both species preferred to mate with conspecific females, but showed no difference in sperm production or expenditure between conspecific and heterospecific females. Females of both species did not prefer conspecific over heterospecific males. Our results suggest that sexual isolation might be mediated by male mate choice in this system and not female choice, suggesting that there is asymmetrical reproductive isolation between the sexes in G. affinis and G. geiseri, but symmetrical species isolation.     

Variation in the reproductive potential of Schistocephalus infected male sticklebacks is associated with 11-ketotestosterone titre

Parasites can impact host reproduction by interfering with host endocrine systems, but the adaptive nature of such effects is disputed. Schistocephalus solidus plerocercoids are parasites of three-spined sticklebacks Gasterosteus aculeatus that are often associated with impaired host reproduction. Here, we relate reproductive behavior and physiology to levels of the androgen 11-ketotestosterone (11KT) in naturally infected and non-infected male sticklebacks from two UK populations. In one population infected males harbored heavy infections and showed uniformly reduced 11KT titres and kidney spiggin (nesting glue protein) content compared to non-infected fish. However in a second population infection levels were more variable and males with smaller infections recorded 11KT and spiggin titres that overlapped those of non-infected fish; among infected males from this population 11KT and kidney spiggin also both correlated negatively with infection severity. Male reproductive behavior correlated closely with 11KT titre in both populations, and infected males with high 11KT levels exhibited normal reproductive behavior. Our results suggest that Schistocephalus infection per se does not block reproductive development in male sticklebacks, and that some male fish may have the ability to breed whilst infected. Our results are not consistent with the hypothesis that Schistocephalus adaptively castrates male hosts via endocrine disruption; rather they support the hypothesis that reproductive disruption is a side effect of the energetic costs of infection.     

Comparison of the organic compounds composition of Hylobius abietis males and females before and after exposure to Beauveria bassiana infection

In the present study, the lipid composition from male and female Hylobius abietis is analyzed pre- and post-infection with Beauveria bassiana. The compounds are analyzed by gas chromatography-mass spectrometry. In the lipids of males and females fatty acids, fatty acid methyl esters, alcohols, amino acids and other organic compounds are identified. The lipids of males and females after B. bassiana exposure contain 31 and 33 organic compounds, respectively, whereas the lipids of non-infected males and females contain 30 and 37 compounds, respectively. The most abundant compounds in infected and non-infected males are fatty acids: C16:0, C18:0, C18:1 and C18:2. Contents of dominant compounds from the lipids of infected and non-infected males are similar to those extracted from the lipids of infected and non-infected females. It should be noted that the total mass of organic compounds is higher in non-infected insects than infected insects.     

Non‐Visual Mate Choice Ability in a Cavefish (Poecilia mexicana) is not Mechanosensory

One of the most important decisions any female has to make is selecting a mate. In many animals, this involves a strong visual component, but in cave‐dwelling fishes, mate preferences have to be exercised in darkness. In a cave‐dwelling Poeciliid fish, the cave molly (Poecilia mexicana), visual mate choice is still possible because they have functional, albeit reduced eyes. In a previous laboratory study, it was documented that females prefer larger males in light and in darkness. In the present study, we investigated the role of the mechanosensory lateral line in this mate preference. We temporarily knocked out the lateral line using antibiotics. We found that even in the functional absence of the lateral line, females showed a clear preference for larger males, indicating that the lateral line is not crucial for mate choice. This points toward a strong role of chemical communication in cave molly mate choice.     

Navigation by Male Crab Spiders <Emphasis Type="Italic">Misumenoides formosipes</Emphasis> (Araneae: Thomisidae): Floral Cues May Aid in Locating Potential Mates

The crab spider Misumenoides formosipes is an ambush predator whose males search for relatively sedentary females within a heterogeneous habitat. Females are receptive to mating immediately after their adult molt and a male biased adult sex ratio together with precopulatory guarding places a premium on male ability to locate females quickly. It is unknown what cues males use to find females; we report here on male movements in association with floral cues. Males in field trials moved towards inflorescences more frequently when both visual and chemical cues were available, than when chemical cues were eliminated. Males in lab trials chose an inflorescence over leaf substrates even in the absence of visual cues. These findings support the hypothesis that M. formosipes males could utilize floral chemistry as a navigational cue in mate searches.     

Shoaling behaviour of sticklebacks infected with the microsporidian parasite, <Emphasis Type="Italic">Glugea anomala</Emphasis>

We compared the shoaling behaviour of three-spined sticklebacks, Gasterosteus aculeatus, infected with the microsporidian, Glugea anomala, to that of non-infected conspecifics. Infected fish lost significantly more weight than non-infected fish during a period of food deprivation, suggesting a metabolic cost to parasitism. In binary shoal choice tests, non-infected test fish showed an association preference for a shoal of non-infected over a shoal of infected conspecifics; infected test fish displayed no preference. Infected fish, however, showed a higher overall tendency to shoal than non-parasitised fish. Furthermore, infected fish occupied front positions within a mixed school. We consider the behavioural differences between infected and uninfected fish in the context of their potential benefits to the fish hosts and the parasites.     

Associative Learning of Color by Males of the Parasitoid Wasp Nasonia vitripennis (Hymenoptera: Pteromalidae)

Males of the parasitoid wasp Nasonia vitripennis showed no innate preference for blue versus yellow or for green versus brown. They learned to associate color with mates, but their ability to do so depended on the color used and the strength of the reward. Specifically, males learned to associate brown or green with a reward of many virgin females. With fewer females, fewer training periods, or mated females as the reward, males still learned a preference for green but not for brown. Males did not learn to associate color with rewards of honey or water. Previous studies of color preference and associative learning in parasitoid wasps have focused almost entirely on females. This is the first demonstration of associative learning in response to visual cues by male parasitoid wasps.     

Male mate choice relies on major histocompatibility complex class I in a sex‐role‐reversed pipefish

Mate choice for compatible genes is often based on genes of the major histocompatibility complex (MHC). Although MHC‐based mate choice is commonly observed in female choice, male mate choice remains elusive. In particular, if males have intense paternal care and are thus the choosing sex, male choice for females with dissimilar MHC can be expected. Here, we investigated whether male mate choice relies on MHC class I genes in the sex‐role reversed pipefish Syngnathus typhle. In a mate choice experiment, we determined the relative importance of visual and olfactory cues by manipulating visibility and olfaction. We found that pipefish males chose females that maximize sequence‐based amino acid distance between MHC class I genotypes in the offspring when olfactory cues were present. Under visual cues, large females were chosen, but in the absence of visual cues, the choice pattern was reversed. The use of sex‐role reversed species thus revealed that sexual selection can lead to the evolution of male mate choice for MHC class I genes.     

Sex recognition in zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

We investigated sex recognition in female zebrafish (Danio rerio) to better understand the underlying sensory mechanisms and identify male secondary sexual traits. Females were simultaneously presented with two fish, a male and a female, in a flow-chamber apparatus, and females’ relative attraction towards males was observed under different conditions. With domesticated fish, females were more attracted to males when presented with both visual and chemosensory cues from stimulus fish. They still discriminated the sexes when only visual cues were provided, but not when white ambient light was changed to yellow, indicating that colour plays a role. Sex discrimination under yellow light was improved when chemosensory cues were also provided. Surprisingly, females’ attraction to males was not more pronounced in the morning when mating occurs. Domesticated females discriminated the sexes when presented with wild-derived, as well as domesticated fish, whereas wild-derived females did not show any biases for domesticated or wild-derived males. Behavioural observations indicated that the wild-derived females were distressed, which explains their lack of attraction to males. In summary, domesticated female zebrafish discriminated the sexes using both visual (body colour) and olfactory cues; however, wild-derived zebrafish were too distressed for behavioural experiments under these laboratory conditions.     

Time spent close to a sexual partner as a measure of female mate preference in a sex-role-reversed population of the blenny<Emphasis Type="Italic"> Salaria pavo</Emphasis> (Risso) (Pisces: Blenniidae)

In most species females do not exhibit conspicuous sexual behaviours and female mate preferences are often measured by means of the time spent close to males. In spite of its widespread use, in only a few studies has this measure been validated as a reliable indicator of female mate preference. Sex-role-reversed species offer a better opportunity to test female preferences, as females usually court males. We tested in a sex-role-reversed population of the blenny Salaria pavo (Risso) if the time spent by females close to males related to the number of courtship displays directed towards males, and whether these measures of female preference reliably predicted matings. Females were simultaneously presented with two males behind a glass partition and the time spent close to each male was measured. We then allowed females to have access to the males' compartments and measured the courtship behaviours performed by females towards each male and recorded with which of the males females spawned. Females spent more time close to the male that subsequently received more courtship displays, and the preferred male had a more developed head crest. However, both measures of female preference failed to predict matings. Females were often attacked by males and probably had, in some occasions, to spawn with the less preferred male. Females that spawned with the previously preferred male had more swollen bellies than females that spawned with the less preferred male, suggesting a male preference towards more ripe females. These results validate the use of "time spent close to a sexual partner" as a measure of female preference in S. pavo. We argue that matings may not always reliably indicate mate preferences, particularly in the less choosy sex.Communicated by R. Serrão Santos     

Female preference for large males in sailfin mollies,<Emphasis Type="Italic"> Poecilia latipinna</Emphasis>: the importance of predation pressure and reproductive status

We examined the effect of predation risk on female association patterns in the live-bearing sailfin molly (Poecilia latipinna). We tested two classes of females, with and without the risk of predation by a green sunfish (Lepomis cyanellus): (1) postpartum females (maintained with males until visibly gravid, then isolated and tested within 24–48 h of dropping a brood); and (2) non-postpartum females (different females, isolated from males for >50 days). When there was no apparent risk of predation, postpartum females showed a significant preference for large over small males, whereas non-postpartum females showed no size preference at all. When there was an apparent risk of predation, postpartum females maintained their preference for larger males and non-postpartum females continued to show no preference for large or small males. These results suggest that reproductive status (receptivity) plays a greater role in mate preference than predation risk. For postpartum females, the cost of not choosing a preferred mate may outweigh the potential cost of predation. Non-postpartum females either do not benefit from mating or are being indecisive about mating and thus are less likely to be choosy whether or not a predator is present.Communicated by I. Schlupp