Sexual selection for male parental care in the sand goby, Pomatoschistus minutus

Male parental care is typically thought to come at a cost to mate attraction and future mating success. However, it has also been hypothesized that paternal care may be under sexual, as well as natural, selection, such that good fathers actually attract more mates. Here we show experimentally that in the sand goby, Pomatoschistus minutus, females prefer to mate with males that provide higher levels of parental care. We manipulated male behavior using (1) different nest sizes and (2) an application of low-O₂ water in the nests, and found that females consistently preferred males with elevated levels of care in dichotomous mate choice tests. This complements our earlier study in which we showed that males increase the amount and quality of care they provide in the presence of females. Our results demonstrate that male care may have evolved as a result of sexual selection rather than natural selection alone, and furthermore, that male care may not necessarily be in conflict with mate attraction.     

The effects of castration, sex ratio and population density on social segregation and habitat use in Soay sheep

We analysed 16 years of census data gathered on the island of Hirta (archipelago of St. Kilda) to investigate the effects of castration, population density, sex ratio, season and group type on habitat use and social segregation of Soay sheep. From 1978 to 1980, 72 male lambs were castrated. We used this experiment to study how a change in reproductive status could affect sociality and habitat choice of these males. Males, females and castrates were all segregated outside the rutting season in autumn. Castrates were the least segregated from females in spring and summer but were most segregated from them during the pre-rut. The more equal the sex ratios, the higher was the degree of social segregation. The three sex classes used similar habitat types, namely, Holcus agrostis, Agrostis festuca and Calluna habitats. Holcus agrostis and Agrostis festuca were top- and second-ranked in female and castrate habitat use, while Holcus agrostis and Calluna were the two top habitat types used by rams. It is unclear why males included Calluna heath habitats, but it cannot be excluded that they might have shifted their use depending on forage availability. A lack in differences in habitat use between castrates and females suggests that body size differences alone cannot be the driving factor for habitat segregation in male and female Soay sheep and that there are reasons other than body size that could motivate reproductive males to use additional habitat types, such as Calluna heath. Although habitat use shifted from one habitat type to the next between low- and high-population-density years and between seasons, there was no clear link between population density and how different groups (male, female or castrate) used these areas. We discuss effects of reproductive status, population density and sex ratio on social segregation and habitat use and suggest that these factors need to be taken into account when investigating causes of sexual segregation in ungulates.     

The impact of behavioral and physiological maternal effects on offspring sex ratio in the common snapping turtle,<Emphasis Type="Italic"> Chelydra serpentina</Emphasis>

Theory suggests that maternal effects are especially important in organisms with environmentally-sensitive sex-determining mechanisms. However, there is no substantive body of empirical evidence to confirm this conjecture. We integrated field and laboratory studies to jointly evaluate the significance of behavioral (nest-site choice) and physiological (yolk hormone allocation) maternal effects on offspring sex ratio in the common snapping turtle (Chelydra serpentina), a species with temperature-dependent sex determination (TSD). Of the 16 microhabitat variables measured, only three (south, east, and total overstory vegetation cover) were significantly correlated with nest temperature: cooler nests were located under more vegetation cover. In turn, these microhabitat predictors of nest temperature, and nest temperature itself, may influence nest sex ratio: shadier, cooler nests were more likely to produce a higher proportion of male offspring than less shady, warmer nests. Analysis of eggs from these same nests incubated in a common garden design in the laboratory revealed that clutch sex ratio was unaffected by levels of yolk estradiol, yolk testosterone, or their interaction. Examination of both behavioral and physiological maternal effects revealed no concordant impact on offspring sex ratio. However, eggs from nests that produced male-biased sex ratios in the field yielded higher proportions of males under constant-temperature conditions in the laboratory. Our study confirms the importance of behavioral maternal effects in nature on offspring sex ratios in species with TSD, while also revealing the potential presence of a predisposition for sex-ratio production underlying TSD in this system.Communicated by S. Krackow     

Female yellowhammers (Emberiza citrinella) prefer yellower males: a laboratory experiment

The importance of male plumage coloration as a signal of male dominance and a cue for female choice in the monogamous yellowhammer, Emberiza citrinella, was tested in two sets of experiments in an indoor aviary. Dominance was tested by introducing two individuals, with no previous experience of each other, in an aviary with food and water. Aggression occurred more often between two males than between a male and a female and more frequently between pairs of males including at least one old male than between two young males. Dominance was not related to male colour in trials between males of the same age class, but it was in trials between an old and a young male, often differing in colour. Thus, age may be a more important determinant of dominance than colour. Female preference for more colourful males was tested by allowing hormone-induced females to choose between a more and a less colourful dummy male. Females spent more time in front of more colourful than drabber males and also more often perched beside colourful males than duller individuals. Although male colour cannot be ruled out as a dominance signal, the results suggest that male colour is primarily used as a signal in mate choice. Female choice may hence be responsible for maintenance of bright plumage in the male yellowhammer.     

Female preferences for multiple attributes in the acoustic signals of the Italian treefrog, <Emphasis Type="Italic">Hyla intermedia</Emphasis>

The advertisement call of frogs and toads is an example of multiple message signal because different acoustic properties encode different kinds of biologically significant information. In the Italian treefrog, Hyla intermedia, pulse rate and frequency have been found to be under stabilizing female preferences and to encode information important for mate recognition, whereas the number of calls per call group have been found to be under directional preferences and, thus, to be important for mate quality assessment. In this study, we investigate preferences for calls that differ simultaneously in frequency, pulse rate, and number of calls per call group, and we ask how these properties interact with each other in influencing female mating decisions. Results of two-choice phonotaxis experiments provide no evidence to support the hypothesis that females process multi-attribute signals in a hierarchical way. In contrast, the pattern of preferences is consistent with the ‘preference function’ hypothesis, that is, with the hypothesis that females rank signals along an ordinal scale of values and choose accordingly. Pulse rate and frequency influence mating preferences more than does the number of calls per call group. The interaction between pulse rate and frequency is not additive but multiplicative: small differences in either pulse rate or frequency that, alone, have no effects on female choice, interact synergistically so that their combination has strong influence on female preferences. A preference repeatability test shows strong among-female differences in preference for multi-attribute signals. We suggest that this result reveals not only a variation in attribute values among females, but also a variation in the way females weight and combine attribute values into a single preference score.     

Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata

In this study, we provide a piece of experimental evidence that immune function is related to dominance and mating success in wild caught male wolf spiders, Hygrolycosa rubrofasciata. In the mating season, H. rubrofasciata males are actively searching for receptive females, and while searching males often engage in agonistic behavior (i.e., agonistic drumming signals, chases, and fights) with each other. The present results demonstrate that dominant males had higher lytic activities in their hemolymph than subordinates. Lytic activity estimates the concentration of antimicrobial peptides with lysozyme-like activity in hemolymph, which have been shown to play an important role in defense against bacteria, viruses, and fungi. Dominants also had higher courtship drumming rates than subordinates. Moreover, winners in mating competition had higher lytic activities than losers, but this was measured nonindependently of dominance status. Among males with mating failure, there was a moderate negative correlation between encapsulation rate and courtship drumming rate, suggesting that low quality males might not be able to bear the immunological costs of courtship behavior. These results suggest that females might gain immunological benefits by preferring vigorous males.     

<Emphasis Type="Italic">Drosophila melanogaster</Emphasis> virgins are more likely to mate with strangers than familiar flies

Recent evidence shows that females of many species can discriminate against males and/or male phenotypes they have mated with previously. However, these studies have not tested whether actual mating is necessary to induce the avoidance behaviour. A preference for strangers may have evolved because it avoids multiple matings with similar genotypes. Alternatively, there may be selection against mating with familiar individuals directly. By choosing its first mate among unfamiliar individuals (which are less likely close relatives than are those encountered early in life), a virgin might disentangle some of the potential benefits of avoiding genetic incompatibility and inbreeding in the offspring from the costs of remating. In this study, we test whether Drosophila melanogaster flies bias their mate choice towards strangers according to previous, non-copulatory, experience. Based on 173 trials over 12 weeks, virgin females presented with two virgin males were 59% more likely to mate with a novel male than the one which she had been housed with for 8 h the day before. Hence we present the first report showing that a dipteran can distinguish between previously encountered and not previously encountered conspecifics.     

Larval host plant origin modifies the adult oviposition preference of the female European grapevine moth <Emphasis Type="Italic">Lobesia botrana</Emphasis>

According to the 'natal habitat preference induction' (NHPI) hypothesis, phytophagous insect females should prefer to lay their eggs on the host species on which they developed as larvae. We tested whether this hypothesis applies to the breeding behaviour of polyphagous European grapevine moth, Lobesia botrana, an important pest in European vineyards. We previously found that different grape cultivars affect several life history traits of the moth. Because the different cultivars of grapes are suspected to provide different plant quality, we tested the NHPI hypothesis by examining oviposition choice of L. botrana among three Vitis vinifera cultivars (Pinot, Chasselas and Chardonnay). In a choice situation, females of L. botrana that had never experienced grapes were able to discriminate between different grape cultivars and preferentially selected Pinot as an oviposition substrate. This 'naive' preference of oviposition could be modified by larval environment: Females raised on grapes as larvae preferred to lay eggs on the cultivar that they had experienced. Furthermore, experience of the host plant during adult emergence could be excluded because when pupae originating from our synthetic diet were exposed to grapes, the emerging adults did not show preference for the cultivar from which they emerged. The NHPI hypothesis that includes the two sub-hypothesis "Hopkins host selection principle" and "chemical legacy" may thus be relevant in this system.     

Divergent evolution of feeding substrate preferences in a phylogenetically young species flock of pupfish (Cyprinodon spp.)

A fundamental question in sympatric speciation is how trophic divergence is achieved. We used an extremely young (<8,000 years) species flock of pupfish (Cyprinodon spp.) from Laguna Chichancanab in south-eastern Mexico to examine divergent evolution of preferences for different feeding substrates. In a test aquarium, we presented four feeding substrates (sand, gravel, a plastic plant, and blank bottom), but no actual food was offered. The four feeding substrates were chosen to mirror the most common substrate types in Laguna Chichancanab. Previous studies demonstrated that benthic food items prevail in the diet of most Cyprinodon species. C. beltrani preferred sand, whereas C. labiosus preferred gravel. F₁ hybrids of both species showed intermediate preferences. C. maya searched for food equally at all substrates. As the test fish were reared under identical laboratory conditions (i.e., in the absence of feeding substrates), the species-specific preferences appear to be genetically fixed, suggesting rapid divergent evolution of feeding behaviors.     

What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich