Foraging ability in the scorpionfly <Emphasis Type="Italic">Panorpa vulgaris</Emphasis>: individual differences and heritability

According to indicator models of sexual selection, mates may obtain indirect, i.e. genetic, benefits from choosing partners indicating high overall genetic quality by honest signals. In the scorpionfly Panorpa vulgaris, both sexes show mating preferences on the basis of the condition of the potential partners. Females prefer males that produce nuptial gifts (i.e. salivary secretions) during copulation, while males invest more nuptial gifts in females of high nutritional status. Both characters, males' ability to produce nuptial gifts and high nutritional status of females, are known to be reliable indicators of foraging ability. Thus, besides possible direct benefits, both sexes might also obtain indirect benefits in terms of “good foraging genes” by their choice and thereby increase the fitness of their offspring. A prerequisite for this possibility is the heritability of the respective trait. In the present study, we estimated the repeatability and the heritability of foraging ability. Our results indicate (1) a significant repeatability of individual foraging efficiencies in males and females and (2) a heritable component of this trait by a significant parent–offspring regression. These findings suggest that genetic benefits in terms of increased offspring foraging ability might contribute to selection for mating preferences in both sexes.     

Indirect fitness benefits are not related to male dominance in a killifish

Recent theoretical and empirical studies have shown that male dominance is often at odds with female mate preference and that indirect (genetic) benefits of mate choice may not be related to male dominance. We tested whether female preference corresponded to male dominance and whether mating with dominant males conveyed benefits to offspring fitness in a small freshwater fish, the African annual killifish Nothobranchius korthausae (Cyprinodontiformes), a species without parental care. The experimental design used controlled for the effect of male age, possibility of sperm and egg depletion, and accounted for a potential that females express their preference through maternal effects by manipulation of egg mass during ovulation. By sequentially mating females with males of known dominance, we found that female N. korthausae showed no mate preference in terms of egg numbers deposited with respect to male dominance or body size and no congruent mate preference to specific males was detected. However, males sired offspring with consistently higher hatching success and the effect was repeatable across individual females. Thus, some males provided females with indirect benefits related to additive genetic quality (“good genes”) and expressed via increased hatching rate, but this benefit was not related to male dominance status or body size.     

Genetic evidence for the “good genes” process of sexual selection

The adaptive nature of female mate choice remains one of the most contentious issues in the study of sexual selection. Here, I provide evidence that mate choice by females of the ovoviviparous cockroach Nauphoeta cinerea influences the rate at which offspring develop and provides both direct and indirect benefits to the female. Males that are more attractive to females produce offspring with shorter development times than less preferred males. Development time is heritable and apparently unconstrained by antagonistic pleiotropy. Male attractiveness and rate of offspring development are genetically correlated. Offspring gain an indirect benefit from their mother's mate choice because, on average, individuals that hatch faster reach sexual maturity more quickly. Females that discriminate among males gain a direct benefit because N. cinerea is ovoviviparous and the time between clutches is decreased by producing offspring with shorter development. In addition to providing evidence for beneficial consequences of mate choice, this study highlights how genetic data provide insights into the process of sexual selection not gained in a purely phenotypic study.Communicated by P. Pamilo     

Male bill colour and age are associated with parental abilities and breeding performance in blackbirds

In monogamous bird species, male parental investment may influence offspring fitness and females may gain advantages through mating with males providing extensive paternal care. However, paternal care is a benefit that can only be assessed indirectly because mate choice precedes paternal activities. Individual quality and age, both signalled by morphological characteristics, may reflect parental abilities. Because they may reflect individual foraging abilities, carotenoid-based colorations have been proposed to honestly signal parental quality. The blackbird (Turdus merula), a socially monogamous species, exhibits biparental care and males show bills that vary from pale yellow to orange due to carotenoid pigments. In this study, we investigated whether male bill colour and age are associated with parental ability. Our results suggest that males with more orange bills and older males are better fathers. Indeed, male visit rate increased with their bill colour index independently of age, and brood condition was higher for adult males, compared to yearlings, independently of bill colour. Overall, the number of fledglings produced was positively influenced by both the age of males and the colour intensity of their bills. Males with more orange bills and adults had a greater number of fledglings and these males also had higher levels of prolactin, a hormone known to promote parental care. This latter finding suggests that prolactin may be the link between carotenoid based colorations and the intensity of paternal effort. Thus, male bill colour seems to honestly reveal male physiological adjustment to paternal activities.     

Parental effects on carotenoid-based plumage coloration in nestling great tits, Parus major

Carotenoid pigments have attracted much interest in behavioural and evolutionary ecology because of their dual function in immune physiology and as color signals. In vertebrates, carotenoids must ultimately be obtained from the diet, and the mechanisms and magnitude of this environmental dependence are central for understanding carotenoid signal functions and evolution. In the present cross-fostering experiment with great tits Parus major, we investigate pre- and postnatal parental effects (egg yolk carotenoids, parental coloration) on nestling size and carotenoid coloration, using HPLC analysis of egg yolk carotenoids, and a reflectance-based measure of ‘chroma’ that reflects the plumage pigment concentration. Both rearing environment and origin influenced offspring size and plumage chroma. Maternal allocation of carotenoids to eggs had a weak positive effect on nestling plumage chroma, whereas we found no prenatal maternal effects (egg size or yolk carotenoid concentration) on size. Nestling plumage chroma was also significantly predicted by the chroma of the rearing father, but not by the color of the rearing mother or either of the original (genetical) parents. Thus, both prenatal maternal effects and postnatal paternal effects influence the carotenoid-based plumage coloration of nestling great tits. Future studies will reveal if parental effects have long-term consequences for plumage development and associated fitness components.     

The importance of biparental care in a precocial,monogamous bird,the bar-headed goose (Anser indicus)

Summary This study presents the first direct evidence of benefit derived from the male's parental effort during rearing of young in a monogamous, precocial bird species. We compared mothers accompanied by a mate (paired mothers) with single mothers, some of which were artificial widows, in a semicaptive flock of bar-headed geese (Anser indicus). Paired mothers were less often displaced by other birds and suffered fewer interruptions of brooding. Their goslings spent more time grazing and survived with a higher probability. Although one beneficial result of the male's parental effort may be the continued availability and survival of his mate, a more direct benefit arises from higher offspring survival between hatching and fledging.     

Female choice on male quantitative traits in lizards — why is it so rare?

Female choice on the basis of male traits has been described in an array of taxa but has rarely been demonstrated in reptiles. In the sand lizard (Lacerta agilis), and possibly in other non-territorial reptiles, a male's contribution to a female's fitness is restricted to his genes. In order to choose males of high genetic quality, females have to trade the fitness gain against the costs of active choice. In a Swedish population of sand lizards, long-lived males sired offspring with higher embryonic survival compared to offspring sired by short-lived males. In spite of this female sand lizards did not mate selectively with older and/or larger males. There appeared to be mo reliable cues to male longevity; age-specific male body size was highly variable. Furthermore, estimates of male nuptial coloration did not covary with ectoparasite load and, hence, females cannot use male coloration as a cue to heritable resistance to pathogenic parasite effects. When cues to male genetic quality are poor, or inaccurate, and males make no parental investment, we predict that female choice will be rare. Sand lizard females mating with many partners lay clutches with higher hatching success. Thus, females may obtain good genes for their young by multiple mating, thereby avoiding costs associated with mate choice.     

Sexual conflict,sexual selection and sperm competition in the spawning decisions of bitterling, <Emphasis Type="Italic">Rhodeus</Emphasis><Emphasis Type="Italic">sericeus</Emphasis>

Sexual selection may operate through either direct selection on preference genes or indirect genetic benefits. However, in many species both direct and indirect selection may operate and can influence female mate and oviposition choice. Adaptations by males for sperm competition can also affect female mate and oviposition choice and can lead to sexual conflict. We investigated the role of direct and indirect benefits and male adaptation to sperm competition for female oviposition decisions in the bitterling, Rhodeus sericeus, a freshwater fish that spawns on the gills of living unionid mussels. Field experiments showed that females chose mussels for spawning on the basis of the number of embryos already in mussels and the body size and/or extent of red coloration of the eyes of territorial males. There was also a significant interaction between these direct and indirect benefits of oviposition choice. Territorial male bitterling avoided leading females to mussels into which rivals had already released sperm. Males also increased their rate of sperm release into mussels in response to sperm release by sneaking males. We discuss the possibility that sexual conflict in oviposition choice occurs in this species.     

Testosterone upregulates lipoprotein status to control sexual attractiveness in a colorful songbird

A salient feature of many secondary sexual characteristics in animals is that their expression is controlled by sex-steroid hormones. However, for only a few types of ornaments do we know the precise molecular mechanism by which androgens like testosterone (T) enhance trait production. We studied the red carotenoid-based beak of male zebra finches (Taeniopygia guttata), which serves as a sexually selected trait and is thought to be T dependent. In previous research, we demonstrated that the extent to which male finches produce lipoproteins in the bloodstream, which bind carotenoid pigments acquired from the diet and transport them to peripheral tissues, regulates the accumulation of carotenoids in the body and beak pigmentation. Here we show that T acts to upregulate lipoprotein production and allows male zebra finches to display flashy, sexually attractive coloration. Levels of circulating T in blood positively and significantly predicted lipoprotein profile (as measured by cholesterol levels), blood carotenoid concentration, and beak color. Exogenous T administration elevated cholesterol and carotenoid status as well as beak redness. Last, experimental inhibition of T (using the anti-androgen flutamide) downregulated lipoprotein production and carotenoid circulation and faded the beak. This androgen- and lipoprotein-mediated system represents one of the more detailed physiological mechanisms underlying the development of a sex-steroid-dependent trait in animals.     

Pre-copulatory sexual selection in the cigarette beetle Lasioderma serricorne

Traditional concepts of sexual selection and sexual conflict make different predictions about the costs and benefits to females of exposure to males with higher mating success. The traditional concepts of sexual selection assume that females benefit from their mate choices, whereas sexual conflict assumes that the females suffer greater costs by mating with males who have greater mating success and thus reduce their fitness. In order to understand how mate choice evolves, it is necessary to estimate the overall effect of mate choice on female fitness. However, relatively few studies have conducted that investigation. In this study, we investigated the direct and indirect effects of mating with attractive males on the fitness of females in the cigarette beetle Lasioderma serricorne. Mating with attractive males increased the number of female offspring but did not affect female longevity. Additionally, we found evidence that attractive males sire highly attractive sons. Thus, mating with an attractive male provides direct and indirect benefits but no fitness cost to female L. serricorne.     

Carotenoid-rich mouth colors influence the conspicuousness of nestling birds

When allocating investment among offspring, parents might maximize their fitness by biasing investment toward offspring with the best direct fitness prospects. The observed preferences of avian parents for carotenoid-rich mouth colors that advertise good condition has been interpreted as support for this hypothesis. However, because these condition-dependent visual signals might also make offspring more visually conspicuous, active parental preferences for carotenoid-rich traits are difficult to distinguish from passive responses to differences in detectability among offspring. Here, we used a visual model to examine how mouth colors influence the visual conspicuousness of nestling house sparrows (Passer domesticus) to parents under a suite of realistic ambient light conditions. We found little evidence that mouths rich in carotenoids provided more conspicuous targets to parents than mouths poor in carotenoids. While other features of mouth color may have evolved to increase conspicuousness, our results suggest that carotenoid-based coloration is not a product of detectability pressures and rather may serve as a signal of nestling quality.     

An experimental test of female choice relative to male structural coloration in eastern bluebirds

Several experimental studies have shown that female birds use ornamental melanin and carotenoid plumage coloration as criteria in mate choice. Whether females choose mates based on natural variation in structural coloration, however, has not been well established. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet (UV)-blue plumage coloration on their head, back, wings, and tail, which is positively correlated with condition, reproductive effort, and reproductive success. We experimentally tested the hypothesis that female eastern bluebirds prefer as mates males that display brighter structural coloration by presenting breeding-condition females with males of variable coloration. We conducted two types of mate-choice experiments. First, females chose between males whose coloration was manipulated within the natural range of variation in the population; feathers were either brightened with violet marker or dulled with black marker. Second, females chose between males with naturally dull or bright plumage coloration. In both manipulated and unmanipulated coloration trials, female choice did not differ significantly from random with respect to structural coloration. We found no support for the hypothesis that the UV–blue coloration of male eastern bluebirds functions as a criterion in female mate choice.     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

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     

Responses of female lizards,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>, to males' chemical cues reflect their mating preference for older males

Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male's age. We hypothesized that female mate-choice preferences may be based on males' pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males' preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.Communicated by W.E. Cooper     

Male care,mate switching,and future reproductive success in a double-brooded passerine

Summary In species with male parental care, polygyny may reduce the parental effort provided by a male, and hence impose a cost on the fitness of his mates, because of decreased growth, survival or health of offspring. I examined a cost of polygyny in the green dart-poison frog, Dendrobates auratus, a species with male parental care in which both male polygyny and mate guarding by females occurs (Summers 1989). All D. auratus males seen carrying tadpoles in a marked area were followed and the pools where they deposited their tadpoles were recorded. Males frequently deposited more than one tadpole in the same pool (in 25% of the observed depositions a male deposited a tadpole in a pool where he had previously deposited at least one other tadpole). Experiments involving manipulation of tadpole densities in pools typically utilized by D. auratus (calabash husks and treeholes) showed that increasing tadpole number had a strong negative effect on both growth rate and survivorship, indicating that polygyny can impose a cost on the fitness of a male's offspring. Hence, females do face a potential cost, in terms of reduced offspring growth and survivorship, when their mates mate polygynously and care for the offspring of other females.     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Communal nursing in the evening bat,Nycticeius humeralis

Summary Nursing observations over two summers involving 76 lactating female evening bats, Nycticeius humeralis, and 128 pups in an attic in northern Missouri indicate that communal nursing occurs rarely until 2 weeks before weaning during which time over 18% of nursing bouts involve nondescendant offspring. The average relatedness among female pairs nursing non-descendant offspring, based on identity-by-descent estimates using allozyme data, was 0.04 (SE=0.12). Mitochondrial DNA d-loop sequence comparisons confirm that at most only 2 of 20 female pairs nursing non-descendant offspring came from the same matriline. Thus, females do not nurse matrilineal kin preferentially despite female natal philopatry. In addition, the average degree of relatedness within a colony (r=0.01, SE =0.03) is too low to provide any indirect benefits from communal nursing. Female error alone is insufficient to explain these observations because females tended to allow female nondescendant young to nurse but excluded nondescendant males, particularly when they had all-male litters. Furthermore, communal nursing bouts did not differ in duration from parental nursing bouts and involved 31 % of all banded females and 24% of all banded pups observed nursing. Communal nursing occurred most frequently when pups began hunting on their own and when lactating females attained their lowest average pre-fed body weight. Mortality during this period was higher for male than female pups, and relative weights implicate starvation as the cause. Time-lapse video records of four families of bats in captivity showed that the number of nursing bouts was proportional to daily weight change. I propose that these results are consistent with both immediate and delayed benefits accruing to females which experience variable hunting success. If a female with extra milk reduced her weight by dumping milk prior to her next foraging trip, she could obtain an immediate energetic benefit and maintain maximum milk production. By restricting such milk donations to nondescendant females she may also increase colony size and thereby enhance her future acquisition of information about foraging and roosting sites.     

No evidence of mate discrimination against males carrying a sex ratio distorter in <Emphasis Type="Italic">Drosophila pseudoobscura</Emphasis>

Selfish genetic elements (SGEs) that spread by manipulating spermatogenesis often have highly deleterious effects on males that carry them. Females that mate with male carriers of SGEs can also suffer significant costs: they receive fewer and poorer-quality sperm, their offspring will inherit the deleterious allele, and the sex ratio of their offspring will be biased towards the more common sex. To counter these costs, females are therefore expected to prefer to mate with males that do not carry sex ratio distorters or other deleterious selfish genetic elements. However, despite the potential costs, there are few examples of female choice against males carrying SGEs. We searched for evidence of a female preference in fruit fly Drosophila pseudoobscura against males carrying a costly meiotic driving X-chromosome Sex Ratio (SR). In a series of five non-competitive mate preference experiments, we find no evidence that females prefer to mate with non-SR males. Our use of five separate experiments, involving more than 800 females, makes it unlikely that this lack of a difference was due to low power or simple chance. We suggest that the lack of female choice against SGE-carrying males may be due to strong selection on SGEs to be indistinguishable from alternative alleles. Furthermore, polyandry, either in direct response to receiving an ejaculate from an SGE-carrying male or carried out indiscriminately when at risk of mating with carriers, may be an alternative response by females to limit the exposure of their offspring to SGEs.     

Carotenoid-based colouration and ultraviolet reflectance of the sexual ornaments of grouse

Among the most familiar sexual signals are red, yellow, and orange sexual traits pigmented by carotenoids. Many birds can detect near-ultraviolet (UV) light, and UV signals can play key roles in mate choice. Grouse (Tetraonidae) exhibit bright carotenoid-dependent sexual ornaments, their supra-orbital combs, which to humans appear orange-red. Combs also reflect in the UV, which is not visible to humans but is likely to be visible to grouse. In male red grouse Lagopus lagopus scoticus, we show that comb UV reflectance decreases with increasing comb size and redness. By removing the epidermis of combs, where carotenoid pigments are, we show that the UV reflectance is a property of the dermis, underneath the red pigmented epidermis. Carotenoid pigmentation of combs acted as a mask to reduce reflectance by the dermis in the range 400–550 nm and in the UV, 300–400 nm. Patagium skin (non-ornamental skin under the wing) also reflects in the UV, but epidermis removal on this bare part tended to reduce UV reflectance, whereas removal of the red epidermis of combs increased UV reflectance. Males in better condition (greater body mass relative to size) had bigger and redder combs, but with less UV. Thus, carotenoid pigments of grouse combs are deposited on a white background with significant UV reflectance, which can influence how the signal is perceived by conspecifics. Carotenoid-based traits exhibit UV reflectance in a number of species, but how UV reflectance and carotenoid pigmentation influence colour remains little known for integumentary ornaments compared to plumage traits. UV vision is not uncommon in birds and other animals, so future studies should investigate how UV reflectance influences the perception of carotenoid-based signals of quality.