Male approach and female avoidance as mechanisms of population discrimination in sagebrush lizards

Reproductive isolation and speciation can result from female choice for particular males. Isolation can also result, however, from male mating preferences or from aggressive encounters which then influence mating decisions. In this study, we use laboratory discrimination trials to study the behavioral mechanisms of population discrimination in sagebrush lizards (Sceloporus graciosus). We specifically ask three questions about population-level discrimination: (1) Does it vary in strength in relation to the geographic distance between the populations? (2) Is it more apparent in inter- or intra-sexual interactions? (3) Does it take the form of attraction or avoidance? We ran 890 trials that tested the ability of male and female sagebrush lizards from one population to discriminate their own population from four other populations. In addition, we utilized both sequential and simultaneous-choice designs, which enabled us to distinguish between attraction and avoidance. We found that most population-level discrimination was exhibited by male lizards preferring to associate with particular types of females, as well as female avoidance of particular types of males. The strength and direction of both discriminations depended on the populations compared and on whether the tests were conducted as sequential- or simultaneous-choice tests, producing a complex relationship between geographic distance and behavioral discrimination. Our results suggest that there are roles for male attraction and female avoidance in population discrimination, reproductive isolation, and speciation.     

Black spots and female association preferences in a sexual/asexual mating complex (Poecilia, Poeciliidae, Teleostei)

We investigated whether female association preferences for males are influenced by black spot disease (BSD), a parasite induced change of the host phenotype. We compared three different species of fish: a gynogenetic hybrid species, Poecilia formosa (amazon molly) and two sexual species (Poecilia latipinna and Poecilia mexicana), which were involved in the natural hybridisation leading to the amazon molly. Contrary to their sexual relatives, asexual amazon mollies significantly avoided images of males infected with black spot disease. We propose that amazon molly females have direct fitness benefits from choosing healthy males. The adaptive significance of the preference for BSD-uninfected males in the asexual amazon molly is yet unclear but may involve avoidance of predation or parasite infection as well as increased sperm availability.     

Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

Indirect genetic benefits of polyandry in a spider with direct costs of mating

The search for the evolutionary explanation of polyandry is increasingly focused on direct and indirect selection on female resistance. In a polyandrous spider Stegodyphus lineatus, males do not provide material benefits and females are resistant to remating. Nevertheless, polyandrous females may obtain indirect genetic benefits that offset the costs associated with multiple mating. We manipulated the opportunity for females to select between different partners and examined the effect of female mating history (mated once, mated twice, or rejected the second male) on offspring body mass, size, condition, and survival under high- and low-food rearing regimens. We found that multiple mating, not female choice, results in increased female offspring body mass and condition. However, these effects were present only in low-food regimen. We did not find any effects of female mating history on male offspring variables. Thus, the benefits of polyandry depend not only on sex, but also on offspring environment. Furthermore, the observed patterns suggest that indirect genetic benefits cannot explain the evolution of female resistance in this system.     

The lemur syndrome unresolved: extreme male reproductive skew in sifakas (<Emphasis Type="Italic">Propithecus verreauxi</Emphasis>), a sexually monomorphic primate with female dominance

The primates of Madagascar (Lemuriformes) are unusual among mammals in that polygynous species lack sexual dimorphism, and females dominate males socially in most species. Moreover, lemur groups are relatively small and characterized by even adult sex ratios despite the fact that one male should be able to exclude other males from the group. One hypothesis to explain this combination of behavioral, morphological, and demographic traits (the “lemur syndrome”) postulates that male–male competition is relaxed and, hence, variance in male reproductive success is low. Reproductive skew theory provides a framework for testing this and several related predictions about lemur social evolution. Specifically, low reproductive skew is also predicted if dominant males or adult females make reproductive concessions to subordinates or if the latter group successfully pursues alternative reproductive tactics. However, suitable data on paternity, demography, and behavior for a conclusive test of these predictions have not been available in the past. In this paper, we show that male reproductive success in ten groups of Verreaux’s sifakas (Propithecus verreauxi) was extremely skewed in favor of the dominant male over 9 years. Our genetic analyses also revealed that more than a third of all groups are effectively harem groups because only one male was unrelated to the resident female(s). In groups with two or more non-natal males, the dominant sired 91% of 33 infants. Together, males pursuing one of several alternative reproductive tactics, such as roaming among several groups or immigrating peacefully, sired only 11% of infants. Thus, female sifakas do not control group composition by offering reproductive opportunities to subordinate males as staying incentives, intrasexual selection is not relaxed, and dominant males prevail in a tug-of-war over subordinate males. Because male reproductive skew in sifakas is even more pronounced than in harem-living anthropoids studied to date, intrasexual selection is clearly not relaxed, and the lemur syndrome is more puzzling than ever.     

Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.Communicated by M. Zuk     

Defensive secretion of a flightless grasshopper: Failure to prevent lizard attack

Summary We tested the role of a grasshopper defensive secretion in deterring lizard predation. Adults, but not young larvae, of the chemically defended lubber grasshopperRomalea guttata (=microptera) froth a volatile secretion when attacked by predators. The lizardAnolis carolinensis failed to strike juvenile lubbers (which lack secretion) in laboratory trials. Survivorship of palatable crickets loaded with secretion offered toA. carolinensis was not significantly different from survivorship of control crickets. In experiments designed to investigate if lizards learn an aversion to the secretion, striking times forSceloporus undulatus fed wax worms coated with secretion were not significantly different over three days of trials. Three primary conclusions are drawn from these data. First, the secretion may not be necessary for lubber protection from lizards. Second, lubber secretion does not appear to deter lizards from attacking or eating prey items. Third, lizards do not appear to develop an aversion to the secretion.     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

Maintenance of variance in sperm transfer rates in a scorpionfly: food availability,genetic basis,and heritability

Variation in traits closely related to fitness is expected to be low. This is because these traits are under directional selection and the best genotype should prevail. However, there have been a number of studies demonstrating the existence of considerable variance in sexually selected traits, which is generally known as the lek paradox. Accordingly, earlier studies found substantial variation in sperm transfer rates in Panorpa vulgaris. Aiming at finding the mechanism that maintains this variation, we analyzed the condition dependence and the narrow sense heritability of sperm transfer rates. Food deprivation in the larval/adult phase caused a decrease in the males’ capability of saliva secretion resulting in shorter copulations and a reduced number of transferred sperm. There was a positive correlation between mean sperm transfer rates and mean body mass. Additionally, intermale variation in sperm transfer rates decreased with increasing food availability. Hence, we suggest that sperm transfer rates in P. vulgaris are influenced by adult feeding history. Heritability analyses of sperm transfer rates did not provide significant results, which is consistent with the general hypothesis that additive genetic variance in traits closely related to fitness is small. Since a trait’s potential to respond to selection is proportional to the amount of contained additive genetic variance, the ascertained small heritability provides a satisfying explanation for the maintenance of substantial variation in sperm transfer rates.     

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.     

Effects of testosterone on male-male competition and male-female interactions in blue tits

In temperate-zone birds, testosterone (T) influences male behavior during the breeding season. The elevation of plasma levels of T to a breeding baseline is necessary for basic reproductive behaviors, but it is still unclear whether variation in T levels above this critical threshold influences the intensity of these behaviors. Such a relationship between T and sexually selected traits is a critical assumption of the immunocompetence handicap hypothesis. We here experimentally elevated T levels in blue-tit males above the natural mean (T-males) during the period of nest building and egg laying, without manipulating hormone levels during chick feeding. T-males neither interacted more often with other males, nor did they respond more aggressively to a dummy intruder, compared to males with control implants. T-males did not guard their social mates more closely, but they were more likely to interact with potential extra-pair mates. Females mated to T-males did not change their behavior during egg laying and the treatment did not significantly affect male and female feeding rates. Despite this, nests of T-males produced larger and heavier fledglings in one study year. Our observations suggest that T levels above the natural mean during the mating period do not increase aggressive or territorial behavior in male blue tits. However, if females perceived T-males as high-quality mating partners, superior offspring development in nests of T-males might be caused by higher maternal investment. Hence, male behaviors involved in mate attraction may have been influenced by T levels above the natural mean.Communicated by R. Gibson     

Male coloration and chick condition in blue-footed booby: a cross-fostering experiment

Sexual selection models suggest that female choice is based on male traits that indicate their genetic or environmental condition, consequently enhancing her reproductive success by direct or indirect benefits. We investigated the relationship between male foot colour and offspring condition in the blue-footed booby, a socially monogamous seabird, with conspicuous and variably condition-dependent coloured feet that are selected by females. In a cross-fostering experiment, we found that chick condition was related to the foot colour of the foster father and, to some extent, to the foot colour of the genetic father; thus overall, the fathers sexual ornamentation (genetic and foster) explained 32% of variance of chick condition. These data suggest that foot colour, a dynamic sexually selected trait, is mostly a signal of parental contribution. In species in which males provide parental care, females may choose mates with higher parental ability. Overall, our data suggest that colourful integuments are honest signals of parental ability.Communicated by J. Graves     

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.     

Sequential male mate choice in a fish,the Pacific blue-eye<Emphasis Type="Italic"> Pseudomugil signifer</Emphasis>

Mate choice is not just a female preoccupation. Under some circumstances, males may also be choosy. However, studies of male mate choice have generally been confined to situations where males can make direct comparisons between potential partners. In contrast, sequential male mate choice has largely been overlooked despite its biologically importance, especially if current investment in mate attraction diminishes a males future mating opportunities. Using the Pacific blue-eye fish Pseudomugil signifer, we show that males are capable of exercising sequential mate choice. When presented sequentially with large and small females, males spent more effort courting the former. However, males did not appear to modify the time spent courting a given female based on the size of the female encountered previously. We suggest that greater attention to the sequential choice problem in males may help illuminate similarities and differences between the sexes when it comes to mating decisions.Communicated by T. Czeschlik     

The effects of larval experience with a complex plant latex on subsequent feeding and oviposition by the cabbage looper moth: Trichoplusia ni (Lepidoptera: Noctuidae)

Summary. Leaf disc choice and oviposition bioassays were used to examine the effects of larval experience with a Hoodia gordonii latex on subsequent behaviors. The latex deterred feeding and oviposition by “naïve” cabbage looper (Trichoplusia ni, Noctuidae) larvae and moths with no previous exposure to the material. “Experienced” insects, reared on a diet with the H. gordonii latex (1000 ppm), exhibited lesser feeding deterrence relative to naïve insects. Experienced female moths actually preferred to lay eggs on treated rather than control leaves. There was no observed transfer of behavioral preferences from experienced parents to their offspring. Our results suggest that moths may be acquiring oviposition preferences from larval feeding experience as described by Hopkins’ host selection principal (HHSP) or through chemical legacy.     

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.     

Do male barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) experience a trade-off between the expression of multiple sexual signals?

Some sexual selection models envisage exaggerated male secondary sexual characters to be costly and therefore reliable indicators of the quality of potential mates to choosy females. If male secondary sexual characters have a natural selection cost, they may be linked to each other by reciprocally constraining relationships that would prevent individual males from increasing their level of multiple signaling. Barn swallow (Hirundo rustica) males have at least two costly signals relevant to socio-sexual interactions: tail length and song. Under the hypothesis that a trade-off exists between male signals, we manipulated the maintenance cost of tail ornaments to test whether this reduced the quantity and quality of song, a condition-dependent, phenotypically plastic signal. Contrary to our expectation, tail elongation had no effect on singing activity and song complexity. However, tail-elongated males produced songs with longer terminal parts ('rattles'). Long rattles are associated with highly competitive social contexts and high circulating levels of testosterone, suggesting that tail-elongated males were more frequently involved in either male-male aggressive or inter-sexual interactions. Therefore, this study shows that males are not displaying multiple signals at the maximum possible level, implying that this system is open to unreliable communication. However, long-term trade-offs between signal expression and viability may prevent males from displaying both signals at higher rates.Communicated by: M. Webster     

Testing a new model of aphid abundance with sedentary and non-sedentary predators

Aphid population dynamics has been thoroughly investigated, especially in tree-dwelling aphids. Among the controls of the aphid rate of increase are the negative effects of antagonists, the positive effects of mutualists, the density-dependence of the aphid dynamics, and the non-stationary quality of plant tissues. Here we present a mechanistic model of aphid growth that considers most of these governing factors using a simple formulation. What is new in this model is that it considers two kinds of antagonists. The first kind is a guild of aphid predator specialists that includes ladybirds (Coleoptera: Coccinellidae), but also species of some families of Hemiptera, Diptera, and Neuroptera. The second kind of antagonists consists of omnivores or generalist predators and in this particular setting is exemplified by the European earwig Forficula auricularia (Dermaptera: Forficulidae). The model developed here compared the effects of these two different kinds of aphid predators, the second one always at the site (sedentary predators) and the first one that arrives in important numbers only once the aphid population has already developed to some degree (non-sedentary predators). Multiple model parameter sets, representing different hypotheses about controls on aphid populations, were evaluated within the Generalised Likelihood Uncertainty Estimation (GLUE) methodology. The model correctly reproduced the experimental data obtained in an organic citrus grove showing the important effect that sedentary predators as earwigs can have on the aphid populations. Low densities of sedentary predators or even low predation rates can have a disproportionate effect on the final aphid density, as they prey on small populations, when the per capita effect on the aphid population is higher. During the main spring peak of aphids the role of non-sedentary predators is secondary, as they track the aphid density rather than control it. However, these non-sedentary predators are important within the proposed model to keep the second autumn peak of aphids at low values.     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.     

Carotenoid-based ornamentation as a dynamic but consistent individual trait

Carotenoid-based ornaments act as signals of quality in many animal species. In contrast to feathers, which are relatively stable structures, carotenoid-pigmented integuments (e.g. bills, lores, tarsi) can change colour rapidly and may better reflect changes in physiological condition. I studied the seasonal variations in plasma carotenoids in red-legged partridges (Alectoris rufa) kept on a constant diet and free of intestinal parasites. Furthermore, I analyzed whether seasonal changes in circulating carotenoids were mirrored by the carotenoid-based coloration of eye rings and bill of this species. Plasma carotenoids showed seasonal variation, with higher levels coinciding with the end of the mating and the start of the laying season. Eye ring pigmentation was related to plasma carotenoid levels, and changes in bill hue (but not changes in UV or red bill chroma) mirrored the variation in plasma carotenoids during the breeding season. Despite the seasonal variation, individual differences in eye ring pigmentation and bill hue, UV and red chroma were consistent throughout the breeding season. Similarly, individual differences in eye ring pigmentation and bill hue and red chroma remained consistent between consecutive years. These results suggest that carotenoid based integumentary colorations act as dynamic traits that accurately reflect the carotenoid-status of individuals, thus reliably indicating consistent differences in individual quality. Furthermore, variability in signal expression appears to have a relevant genetic/phenotypic basis independently of environmental conditions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.