Can aposematic signals evolve by gradual change?

Aposematic species, which signal conspicuously of their unprofitability to predators, have puzzled evolutionary biologists for over a century1,2. Although conspicuousness of unpalatable prey improves avoidance learning by predators3,4,5, it also involves an evolutionary paradox: with increasing detectability4,6,7,8 the deviant aposematic prey would suffer high predation initially from naive predators. Here we test a neglected idea7,8,9,10,11 that aposematic coloration may evolve by gradual change rather than by major mutations. Weak signals did not suffer high initial predation, but predators (great tits, Parus major) did not learn to separate them from cryptic palatable prey. Furthermore, …

Lek centre attracts black grouse females

In several lek mating systems, centrally located males enjoy higher mating success than peripheral males. The mechanism behind this pattern, however, has been controversial and a rigorous test of the different alternatives is missing. Here we report that in black grouse Tetrao tetrix , a lekking bird species, central males achieved many more copulations than males in the periphery. Generally, central territories were smaller than peripheral ones, and males were thus more clumped in the centre. A choice experiment in an aviary provided experimental support for females being attracted to densely clustered males, so that males in larger clusters have higher expected average mating success. Thu…

Can ultraviolet cues function as aposematic signals?

The fact that birds are sensitive to ultraviolet light (UV, 320–400 nm) has been largely ignored by previous studies of aposematism. Therefore, in the present article we investigated whether great tits preferred ultraviolet-reflecting colors compared to colors without UV reflection and whether UV cues alone could function as aposematic signals. We were able to manipulate prey visibility in UV light by changing the UV reflectance of prey items as well as altering the lighting conditions. In order to perform a preference experiment we used three pairs of colors (green UV vs. green, gray UV vs. gray, yellow UV vs. yellow) on a black background. The birds ate both UV types equally for all three…

Male coloration and species recognition in sympatric flycatchers

Currently favoured views for explaining ornaments in males emphasize female preference such that females benefit from increased offspring production, good genes of the offspring, or the attractiveness of sons. Results from long-term studies in the pied flycatcher Ficedula hypoleuca suggest that factors associated with species recognition may also be important for male coloration. In sympatry the collared flycatcher F. albicollis is dominant in competition for nesting sites over the pied flycatcher. Bright pied flycatcher males resemble collared flycatcher males and suffer from interspecific interference, whereas dull and female-like males can acquire nesting sites close to those of the coll…

Food supplementation reveals constraints and adaptability of egg quality in the magpie Pica pica

Differences in the deposition of limited maternal resources to eggs can reflect the optimal allocation to manipulate offspring phenotype, or constraints caused by maternal condition. We examined multiple maternal substances transferred to eggs in the magpie Pica pica to test the hypothesis that certain substances represent constraint and some optimal allocation. We did this by supplementary feeding magpies prior to egg-laying and then comparing the effect of food on maternal substances in conjunction with laying order relative to a control group. Certain substances such as carotenoids, immunoglobulins and avidin responded positively to food supplementation, whereas others, such as testoste…

Restrictive mate choice criteria cause age-specific inbreeding in female black grouse, Tetrao tetrix

Inbreeding is generally rare in large, natural populations yet mate choice often appears to be random with respect to kinship. This suggests that the risks of inbreeding may be small because passive mechanisms of inbreeding avoidance, for example dispersal, are effective at lowering inbreeding risk. Previous theoretical and empirical studies have assumed that the risks of inbreeding are constant over an individual’s life span, but in the lek-breeding black grouse, inbreeding increases with female age. To determine whether inbreeding avoidance mechanisms are also age dependent, we generated four null models of random mate choice ranging from complete randomness to more biologically realistic…

The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix)

We describe the results of two studies of parasitic infection in the black grouse (Tetrao tetrix). The first deals with our own observations of lekking black grouse in which the parasite levels of two blood parasites, the protozoan Leucocytozoon lovati and microfilaria, probably produced by a nematode worm Splendidofilaria tuvensis, were scored. We also obtained measures of age, survival, number of copulations, body mass and length of the ornamental tail feathers (the lyre) of the lekking males. The second study analysed the data from Lund (1954) which involved eight gut parasites obtained from birds which were killed. In the first study we found higher levels of infection of Leucocytozoon …

Extra-pair paternity and male characteristics in the pied flycatcher

The pied flycatcher (Ficedula hypoleuca) is sexually dichromatic with extreme variation in male plumage coloration. The benefit for males of having black plumage is controversial, and few studies have found evidence for a sexual selection benefit of being black rather than brown. However, blacker males may be better able to achieve extra-pair fertilizations (EPFs), which may be an important component of sexual selection. We studied the role of EPFs in sexual selection in the pied flycatcher by establishing a set-up where two males with different back coloration (blacker vs browner) bred simultaneously near each other. DNA fingerprinting analysis revealed that 11% of offspring resulted from …

Paternity, copulation disturbance and female choice in lekking black grouse

Female copulation behaviour in the black grouse, Tetrao tetrix, was studied by detailed observations of individually marked birds together with DNA fingerprinting analyses for paternity assessment. For each breeding occasion females typically mated only once with one male, and did not mate outside the lek arena. They re-mated with the same or another male usually only if the initial copulation was disturbed and was probably unsuccessful in sperm transfer. The willingness of females to copulate only once with one male suggests that multiple mating with several males incurs a cost that more than outweighs any possible fertility or sperm competition benefits. Attempts by neighbouring males to …

Determinants of the Mating Success of Polyterritorial Pied Flycatcher Males

Much attention has been paid to the polyterritorial mating system of some passerine birds. Here we report how a male's mating success is related to the behavioral traits of polyterritorial pied flycatcher (Ficedula hypoleuca) males. We found no evidence that the timing of polyterritoriality in relation to egg laying in the primary nest or the singing behavior of males have any influence on mating success. However, results show clearly that male mating success was improved with an increase in the distance between territories up to a distance of about 200–300 m whereupon there was no further enhancement of mating success. This finding is crucial for both the deception hypothesis and female-fe…

Why Do Black Grouse Males Perform on Lek Sites outside the Breeding Season?

Studies of sexual selection have focused mainly on explanations of how male characteristics influence mating success during the breeding season. By contrast, few studies have investigated how behaviours prior to the breeding season influence mate choice d

Do palatable prey benefit from aposematic neighbors?

Synergistic selection can promote the evolution of aposematism. This requires, however, that palatable prey without signals do not benefit from their aposematic neighbours. In order to test this re...

Prehatching maternal investment and offspring immunity in the pied flycatcher (Ficedula hypoleuca).

Maternal investment in offspring immunity via egg quality may be an adaptive evolutionary strategy shaped by natural selection. We investigated how maternal investment in eggs can influence offspring immunity by conducting two experiments. First, we manipulated foraging performance of the mothers before egg laying by attaching a small weight to their back feathers. During the nestling period, we investigated offspring total antibody production at the age of 7 days and after antibody challenge, and conducted a partial cross-fostering design to separate the effects of the experiment and rearing-related variation on offspring immunity. In a separate experiment, partial cross-fostering with ant…

Assortative mating and female clutch investment in black grouse

Variation in female behaviour has only recently received attention in studies of sexual selection. It has been suggested that females may invest differentially in their offspring in relation to the quality of their mate. This may lead to females that mate with high-quality and/or attractive males laying larger clutches. Females may also differ in their ability to choose between males. For example, females in good physical condition may make better choices. If physical condition and clutch size are positively correlated, this hypothesis could also produce a relationship between male attractiveness and female clutch size. We found, in lekking black grouse, Tetrao tetrix, that females mated to…

Imperfect Batesian mimicry—the effects of the frequency and the distastefulness of the model

Batesian mimicry is the resemblance between unpalatable models and palatable mimics. The widely accepted idea is that the frequency and the unprofitability of the model are crucial for the introduction of a Batesian mimic into the prey population. However, experimental evidence is limited and furthermore, previous studies have considered mainly perfect mimicry (automimicry). We investigated imperfect Batesian mimicry by varying the frequency of an aposematic model at two levels of distastefulness. The predator encountered prey in a random order, one prey item at a time. The prey were thus presented realistically in a sequential way. Great tits (Parus major) were used as predators. This expe…

Female-female aggression and male polyterritoriality in the pied flycatcher

The search cost in mate choice of the pied flycatcher

Sexual selection in a wolf spider: Male drumming activity, body size, and viability

Females are often believed to actively choose highly ornamented males (males with extravagant morphological signals or intense sexual display), and ornaments should be honest signals of male viability. However, this belief is relying only on some pieces of empirical evidence from birds. Our study reports active female choice on sexual display that indicates male viability in spiders. We established trials in which we studied female choice in relation to male courtship drumming activity and body size. Females chose the most actively drumming males as mating partners, but the body size of the males did not seem to be selected. Male drumming activity turned out to be a good predictor of male v…

Genetic Component of Morphological Differentiation in Coal Tits Under Competitive Release

Mate sampling behaviour of black grouse females (Tetrao tetrix)

We studied female mate sampling behaviour in lekking black grouse (Tetrao tetrix). Females mainly visited males occupying territories in the centre of the lek with relatively large territories. They were also more likely to visit males that had high attendance. The same factors were also correlated with male mating success. A multiple regression model including these factors explained more of the variance in female visits per male (53%) than in mating success (33%). The pattern of female sampling conformed with a pool comparison (best-of-n) tactic. Such a tactic is expected if the costs of sampling are low. Females of high body mass visited more males than lighter females, however, which in…

Large-scale drainage and breeding success in boreal forest grouse

Summary 1 The breeding success of Finnish grouse has been in decline for decades. While it has been shown that fragmentation and modern forestry practices such as clear-cutting affect the viability of grouse populations, little is known about effects of large-scale drainage. The drainage network in Finland has increased dramatically during the past decades. By 1988, 6 million ha of bog ecosystems had been drained. This is likely to have had a profound direct and indirect effects on grouse productivity. Because ditches persist in time, large-scale drainage may therefore have strong potential for affecting the long-term breeding success of three forest grouse species in Finland. 2 Using a mix…

Can female preference explain sexual dichromatism in the pied flycatcher, Ficedula hypoleuca?

How important female choice is for the evolution of male secondary sexual characteristics is controversial. Two field and one laboratory experiment, using the pied flycatcher, were performed to test the female choice aspect of sexual selection. In addition, non-manipulative data from 5 years are presented. The observational data suggest a slight preference for dark males by females but in field experiments in which males had territories at random sites (i.e. they did not choose a territory) or the colour of concurrently arriving males was altered, there was no preference for darker ones. Similarly, oestradiol-treated females did not prefer black or brown males in the laboratory. Thus, there…

GENETIC AND ENVIRONMENTAL VARIATION IN ANTIBODY AND T-CELL MEDIATED RESPONSES IN THE GREAT TIT

Host parasite coevolution assumes pathogen specific genetic variation in host immune defense. Also, if immune function plays a role in the evolution of life history, allocation to immune function should be heritable. We conducted a cross-fostering experiment to test the relative importance of genetic and environmental sources of variation in T-cell mediated inflammatory response and antigen specific antibody responses in the great tits Parus major. Cell mediated response was measured during the nestling period and antibody response against two novel antigens was measured in two-month-old juveniles raised in a laboratory. We found no effect of nest of origin, but a strong effect of rearing e…

Low levels of relatedness on black grouse leks despite male philopatry

In lekking species, males cluster on specific areas for display (the leks) and females generally prefer to copulate with males on large aggregations. The maintenance of leks in which only a few males reproduce might be explained if subordinate males gain indirect fitness benefits. By joining a lek on which relatives are displaying, subordinates might attract more females to the lek thereby increasing the mating opportunities of their kin. In black grouse, a genetic structure among leks has previously been found suggesting that relatives could display together. Using 11 microsatellite loci, we extended this result by testing for the presence of kin structures in nine black grouse leks (101 m…

Limited indirect fitness benefits of male group membership in a lekking species

In group living species, individuals may gain the indirect fitness benefits characterizing kin selection when groups contain close relatives. However, tests of kin selection have primarily focused on cooperatively breeding and eusocial species, whereas its importance in other forms of group living remains to be fully understood. Lekking is a form of grouping where males display on small aggregated territories, which females then visit to mate. As females prefer larger aggregations, territorial males might gain indirect fitness benefits if their presence increases the fitness of close relatives. Previous studies have tested specific predictions of kin selection models using measures such as …

Founder population size and number of source populations enhance colonization success in waterstriders.

Understanding the factors that underlie colonization success is crucial both for ecological theory and conservation practices. The most effective way to assess colonization ability is to introduce experimentally different sets of individuals in empty patches of suitable habitat and to monitor the outcome. We translocated mated female waterstriders, Aquarius najas, into 90 streams that were not currently inhabited by the species. We manipulated sizes of propagules (from 2 to 16 mated females) and numbers of origin populations (one or two). Three origin populations were genetically different from each other, but they were less than 150 km from the streams of translocation. The results demonst…

Overwintering survival in relation to body mass in a field population of the wolf spider ( Hygrolycosa rubrofasciata )

Body size is often considered to be an important trait affecting individual fitness. In arthropods, females commonly benefit from larger size directly through increased fecundity (Roff, 1992), and males through increased mating success (Andersson, 1994). It has also been suggested that larger individuals may in general have a better survival than smaller individuals (Calder, 1983; Peters, 1983). From this suggestion it may be predicted that during stressful environmental conditions larger individuals should do better than smaller individuals.

Copying the Mate Choice of Others? Observations On Female Black Grouse

Abstract It has been suggested that the non-random mating often observed in lekking species is a consequence of either male-male competition or active female mate choice. Here we show that the highly skewed mating distributions observed in a black grouse lek in three years were indeed different from random expectations. We suggest that females copying the mate choice of others enhance this skew. Observations in favour of copying are: females pay multiple visits to the lek during several days; females arrive and move in bands which makes it possible to observe the visits to male territories and matings of other females; in the main lek in the study area, males often mated in sequence indica…

Context-dependent effects of tail-ornament damage on mating success in black grouse

Evolution of black grouse leks: female preferences benefit males in larger leks

Joint brood guarding in parent bugs — an experiment on defence against predation

Females of Elasmucha grisea defend their eggs and small nymphs against invertebrate predators. Females sometimes guard their clutches side by side on the same birch leaf. We studied benefits of this joint guarding both in the field and in the laboratory. We found that adjacent females had significantly larger clutches than solitary females. In the laboratory, we studied the effectiveness of joint versus single defence against ant (Formica uralensis) predators. We established female pairs from initially singly guarding females by cutting off pieces of leaves with egg clutches and pasting them beside another female guarding her clutch. In the control group the females with their clutches were…

Microhabitat selection and audible sexual signalling in the wolf spider Hygrolycosa rubrofasciata (Araneae, Lycosidae)

We studied the microhabitat selection and male sexual signalling behaviour in the wolf spider Hygrolycosa rubrofasciata (Ohlert). Males strike dry leaves with their abdomen, producing an audible sexual drumming signal, and females use this signal to choose mating partners. In the field we followed male drum- ming rate and microhabitat selection using both the mark-recapture method and direct observations. In the laboratory we conducted an experiment on male micro- habitat and drumming substrate selection. We found that in the field males were not distributed randomly among the habitat; fewer males were found in areas that had high sedge cover, low elevation, and low dry leaf cover. In the l…

Female choice for male drumming in the wolf spiderHygrolycosa rubrofasciata

Mate preferences in invertebrates have usually been studied with simultaneous choice exper- iments alone, which allows eVective detection of any preferences but does not tell much about the strength of inter-sexual selection. Under natural conditions females frequently have to rely on sequential choice, and choosy females may incur opportunity and direct costs such as loss of time when they reject a male. Female preference in the wolf spider Hygrolycosa rubrofasciata for two components of male courtship signalling, rate and volume, was investigated. Both of these characteristics were tested with a sequential choice set-up and the eVect of volume also with a simultaneous choice method. Femal…

Alarm calling by individual willow tits, Parus montanus

Abstract Alarm responses of wild-captured individual willow tits to model sparrowhawks, Accipiter nisus, flying over a test chamber were studied. Tits did not usually give alarm calls if the apparent predator was passing nearby (at a height of 10 m), while over half of the individuals responded by alarm calling for a more distant predator (40 m). This suggests that alarm calling involves a risk to the caller. Second, there was individual variation in the responses, with older males giving the call more frequently than females or young males. The greater responsiveness of adult males may indicate that different individuals in a flock gain different benefits by warning the others. The tendenc…

Heritabilities and paradigm shifts

Evolution of foraging ecology in Fennoscandian tits ( Parus spp.)

Species belonging to the same genus exploit similar resources in a similar manner. Despite their general similarities in behaviour, they differ in morphology, and these differences are often interpreted as adaptations for use of different resources. Here we report on the relation between morphology, habitat selection and feeding ecology in seven tit species (Parus spp.) in Fennoscandia. A problem previously disregarded in most ecomorphological studies is that members of a guild with closely related species share their common ancestry. This historical legacy causes similarity in ecology and morphology. We have used phylogenetic information with proper comparative methods to reveal the evolut…

Arena birds

Prenatal androgen exposure modulates cellular and humoral immune function of black-headed gull chicks

Avian eggs contain considerable amounts of maternal yolk androgens, which have been shown to beneficially influence the physiology and behaviour of the chick. As androgens may suppress immune functions, they may also entail costs for the chick. This is particularly relevant for colonial species, such as the black-headed gull (Larus ridibundus), in which the aggregation of large numbers of birds during the breeding season enhances the risk of infectious diseases for the hatching chick.To test the effect of maternal yolk androgens on the chick's immune function, we experimentally manipulated, in a field study, yolk androgen levels within the physiological range by in ovo injection of either a…

Life-history differences in age-dependent expressions of multiple ornaments and behaviors in a lekking bird

Age is a major factor explaining variation in life-history traits among individuals with typical patterns of increasing trait values early in life, maximum trait expression, and senescence. However, age-dependent variation in the expressions of sexually selected traits has received less attention, although such variation underpins differences in male competitive abilities and female preference, which are central to sexual selection. In contrast to previous studies focusing on single traits, we used repeated measures of seven sexually selected morphological and behavioral traits in male black grouse (Tetrao tetrix) to quantify the effects of age and life span on their expressions and quantif…

Polyterritoriality and deception

Testosterone and male mating success on the black grouse leks

On black grouse (Tetrao tetrix) leks females prefer dominant, viable males that have managed to acquire relatively central territories. The immunocompetence hypothesis predicts that, because high levels of testosterone are costly to the immune system, male sexual traits that are controlled by testosterone are likely to serve as reliable indicators of male health. Indeed, testosterone concentrations of black grouse males were highly variable, and strongly correlated with male mating success. This is related to the fact that males with high testosterone levels had most central territories. However, the association of testostestorone level with male mating success was not solely related to cen…

Responses of Male and Female Black Grouse to Male Vocal Display

The evolution of leks may be explained by several hypotheses. The ‘female preference’ hypothesis, which states that females favour males that have aggregated, has recently gained some empirical support. Low-quality, unattractive males may, however, settle near attractive males, as predicted by the ‘hotshot’ hypothesis. We tested whether black grouse Tetrao tetrix females use auditory cues to find the preferred leks, and whether males respond to vocal display emitted on leks. We conducted a playback experiment with male vocal display (rookooing) on leks, where the visiting females and displaying males were counted. The number of males tended to increase more on playback leks. Specifically, t…

Multiple benefits of gregariousness cover detectability costs in aposematic aggregations

Understanding the early evolution of aposematic (warning) coloration has been a challenge for scientists, as a new conspicuous morph in a population of cryptic insects would have a high predation risk and would probably die out before local predators learnt to avoid it1,2,3,4. Fisher5 presented the idea of aggregation benefit through the survival of related individuals; however, his theory has been strongly debated6,7,8 as the mechanisms that favour grouping have never been explored experimentally with the incorporation of detectability costs. Here we create a comprehensive ‘novel world’ experiment with the great tit (Parus major) as a predator to explore simultaneously the predation-relate…

A trade-off between sexual signalling and immune function in a natural population of the drumming wolf spider Hygrolycosa rubrofasciata.

The field of ecological immunology is ultimately seeking to address the question ‘Why is there variation in immune function?’ Here, we provide experimental evidence that costs of ubiquitous sexual signals are a significant source of variation in immune function. In the mating season, males of the wolf spider Hygrolycosa rubrofasciata drum against dry leaves while wandering around the habitat searching for receptive females. According to a previous study, the male metabolic rate during the drumming increases 22-fold compared to the resting metabolic rate. In the present study, we examined whether investment in costly courtship drumming decreases male immune function in a wild population of H…

Female choice in the kestrel under different availability of mating options

Abstract Mate choice in relation to its fitness consequences on females was studied in kestrels, Falco tinnunculus , in western Finland. The breeding season is short with rapidly declining prospects of successful breeding, suggesting that females should make their mating decision quickly. On average, males arrived a few days before females, but late males arrived much later than early females. Pairs were usually formed within 2 days of male arrival, and only early males were available to early females. Options for female choice differed considerably in years of different kestrel density. In a good vole year, 1988, females had many males to choose from, but in poor vole years, 1989 and 1990,…

Maternal antibody transmission and breeding densities in the Black-headed Gull Larus ridibundus

1. The offspring of avian species, especially those of colonial breeders, are exposed to a number of pathogens immediately after birth. The chick's immune system is, at that early stage still immature and inefficient. As a consequence, diseases can have a strong impact on chick survival.2. The ability of mothers to transmit passive immunity in terms of antibodies of their own acquired immunity to their chicks is probably an essential pathway to enhance the chick survival. Since the production of antibodies is costly, females are expected to adjust the transmission of passive immunity to the local disease environment.3. We found that in Black-headed Gulls (Larus ridibundus L.) yolk antibody …

Restrictive mating by females on black grouse leks

In bird species with pair bonds, extra-pair matings could allow females to choose genetically superior males. This is not needed in lekking species because female choice is not constrained by pairing opportunities. However, polyandry has been reported in most lekking species studied so far. Using 12 microsatellite loci, we determined the paternity of 135 broods of black grouse sampled between 2001 and 2005 (970 hatchlings and 811 adult birds genotyped). The paternity assignments were combined to lek observations to investigate the mating behaviour of black grouse females. About 10% of the matings seemed to take place with males displaying solitarily. Forty per cent of the copulations betwee…

Habitat Differences and Variability in the Lek Mating System of Black Grouse

Spatially fixed territoriality has been regarded to be a general feature in lek mating systems. In the absence of territories, however, the nature of leks may change remarkably. We compared leks of black grouse in areas, where territoriality was governed by habitat characteristics. On ice-covered lakes, where the lekking ground was homogeneous and offered no landmarks by which the displaying males would be able to recognize territory boundaries, the male aggregations were mobile. 'Classical' leks on bogs, however, were stable. It also appeared that on mobile leks, as compared to stable leks, the opportunities for females to choose their mates were reduced. The distribution of mating success…

Strong antiapostatic selection against novel rare aposematic prey

The evolution of aposematism, a phenomenon where prey species conspicuously advertise their unprofitability to predators, is puzzling. How did conspicuousness evolve, if it simultaneously increased the likelihood of an inexperienced predator to detect the prey and presumably kill it? Antiapostatic selection, where rare prey is predated relatively more often, is considered as another major difficulty for aposematism to evolve. However, the risk of being conspicuous in low frequencies has not been experimentally tested. We designed an experiment to test how frequency (4%, 12%, 32%) of conspicuous aposematic prey and its dispersion type (solitary vs. aggregated) affect an initial predation ri…

Sexual selection in the wolf spider Hygrolycosa rubrofasciata: female preference for drum duration and pulse rate

The unusual form of sexual signaling, the drumming produced by the wolf spider Hygrolycosa rubrofasciata, allows exceptionally detailed studies of female preference patterns against signal characteristics. It is easy to manipulate the signals and to use large numbers of females in playback experiments. Males of H. rubrofasciata produce drums by striking their abdomen against dry leaves on the ground. Drums travel not only as substrate-borne vibrations, but also as airborne acoustic signals. Females respond sooner to drums transferred as substrate borne, but the mode of signal transfer has no effect on female preference for different types of drums. We investigated the effects of two key com…

Effects of novelty and gregariousness in survival of aposematic prey

We examined the reactions of captive wild great tits to novel unpalatable prey with (1) a traditional aposematic signal (black and yellow) (2) a novel signal (light pink), and (3) a control signal (brown). Prey were offered either singly or in groups to see whether novel signals with fewer possibilities for synergistic benefits are more dependent on grouping than are the traditional signals. Indeed, responses of birds toward unpalatable prey depended significantly on spatial distribution of prey (grouping versus solitary) and die type of die signal. Birds avoided more the traditional black and yellow signals than novel pink signals in both experimental set ups, but both of diese prey items …

Singing rate and female attraction in the pied flycatcher: an experiment

«Attirer les femelles» est une des principales fonctions du chant des oiseaux. Beaucoup d'etudes ont mis en evidence l'effet de variations du repertoire sonore sur le succes d'accouplement mais peu d'entre elles sont consacrees a l'intensite du chant. Dans une telle optique, cette etude examine les effets de manipulations experimentales de chants sur la selection du partenaire par les femelles

Good-genes effects in sexual selection

The magnitude of the effect of good genes as a viability benefit accruing to choosy females remains a controversial theoretical and empirical issue. We collected all available data from the literature to estimate the magnitude of good-genes viability effects, while adjusting for sample size. The average correlation coefficient between male traits and offspring survival in 22 studies was 0.122, which differed highly significantly from zero. This implies that male characters chosen by females reveal on average 1.5% of the variance in viability. The studies demonstrated considerable heterogeneity in effect size; some of this heterogeneity could be accounted for by differences among taxa (birds…

Acoustic signalling in a wolf spider: can signal characteristics predict male quality?

While there has been considerable interest in female choice for male sexual signals, there have been few studies of the underlying information that different aspects of the signal calls convey. Such studies, however, are essential to understand the significance of signals as honest handicaps, arbitrary Fisherian traits and/or in species recognition. We studied the somewhat exceptional system of audible drumming in the wolf spider Hygrolycosa rubrofasciata. We estimated the repeatabilities of signal components, the levels of between-male variance, the symmetry of the signal, the correlations between different aspects of drumming and their correlations with body weight. While in other taxa th…

Do female leaf beetles Galerucella nymphaeae choose their mates and does it matter?

The role of active female choice in sexual selection is frequently difficult to ascertain, and this is particularly the case for many insect species. Also, it is uncertain whether choosing between males would affect offspring viability. We designed an experiment to investigate the presence of female choice in a Coleoptera species (Galerucella nymphaeae). We also estimated whether mate choice would have any effect on offspring performance. Females were first placed with two males in a test arena to see which of the males copulated with the virgin female, and how quickly. Subsequently the loser male was offered a new virgin female to test for any change in latency time until mating. The two-m…

Plumage colour and male-male interactions in the pied flycatcher

Abstract Abstract. The influence of male colour on male-male interactions in the pied flycatcher, Ficedula hypoleuca , was studied using several types of field experiments. This species exhibits delayed plumage maturation, each male becoming more conspicuously black and white from the first to the second breeding summer. However, individual males vary even more, some old males being as brown as females. When territorial males were presented with a choice between brown and black male intruders (or brown males painted black) they directed more aggression towards the black males. Brown plumage did not help males obtain nestboxes close to other males. If anything browner males were at a disadva…

Short- and long-term population dynamical consequences of asymmetric climate change in black grouse

Temporal asymmetry in patterns of regional climate change may jeopardize the match between the proximate and ultimate cues of the timing of breeding. The consequences on short- and long-term population dynamics and trends as well as the underlying mechanisms are, however, often unknown. Using long-term data from Finland, we demonstrate that black grouse ( Tetrao tetrix ) have responded to spring warming by advancing both egg-laying and hatching. However, early summer (the time of hatching) has not advanced, and chicks have to face colder post-hatching conditions. Demonstrating that these conditions are critical to post-hatching survival, we show that chicks are increasingly suffering highe…

Energetic costs of size and sexual signalling in a wolf spider

A prerequisite for honest handicaps is that there are significant condition–dependent costs in the expression of sexual traits. In the wolf spider Hygrolycosa rubrofasciata (Ohlert), sexual signalling (drumming) is costly in terms of increased mortality. Here we investigated whether this mortality may be caused by increased energy expenditure. During sexual signalling, metabolic rate was 22 times higher than at rest and four times higher than when males were actively moving. Metabolic rate per unit mass was positively related to absolute body mass during sexual signalling but not during other activities. This positive relationship is novel to any studies of metabolic rates. Indeed, it seems…

Age-dependent inbreeding risk and offspring fitness costs in female black grouse

Dispersal is an important mechanism used to avoid inbreeding. However, dispersal may only be effective for part of an individual's lifespan since, post-dispersal individuals that breed over multiple reproductive events may risk mating with kin of the philopatric sex as they age. We tested this hypothesis in black grouse Tetrao tetrix , and show that yearling females never mated with close relatives whereas older females did. However, matings were not with direct kin suggesting that short-distance dispersal to sites containing kin and subsequent overlap of reproductive lifespans between males and females were causing this pattern. Chick mass was lower when kinship was high, suggesting impor…

Sexual advertisement and immune function in an arachnid species (Lycosidae)

A simple version of the immunocompetence handicap hypothesizes that through condition-dependence, the size of the sexual trait may be positively related to immune function at the population level. In the present study, we investigated the relationship between sexual advertisement and immune function in a natural population of male wolf spiders, Hygrolycosa rubrofasciata (Araneae: Lycosidae). Males of H. rubrofasciata have a costly and condition-dependent acoustic signal, courtship drumming. In the mating season, males drum against dry leaves while wandering around the habitat searching for receptive females. Males increase their mating success by increasing their drumming rate and mobility.…

Inbreeding depression and male fitness in black grouse

The male lifetime lekking performance was studied, and related to inbreeding-outbreeding in a wild population of black grouse (Tetrao tetrix) in central Finland between 1989 and 1995. Inbreeding was measured as the mean heterozygosity and mean d(2) of 15 microsatellite loci. We found a significantly positive relationship between mean d(2) and lifetime copulation success (LCS), while the relationship between heterozygosity and LCS was close to significant. We also found that males that never obtained a lek territory had significantly lower mean heterozygosity than males that were observed on a territory at least during one mating season in their life. Furthermore, among males that were succe…

SIGNAL RELIABILITY COMPROMISED BY GENOTYPE-BY-ENVIRONMENT INTERACTION AND POTENTIAL MECHANISMS FOR ITS PRESERVATION

Sexual selection based on signaling requires that signals used by females in mate choice are reliable indicators of a male's heritable total fitness. A signal and the preference for it are expected to be heritable, resulting in the maintenance of genetic covariance between these two traits. However, a recent article has proposed that signals may quickly become unreliable in the presence of both environmental variation and genotype-by-environment interaction (G x E) with crossing reaction norms, potentially compromising the mechanisms of sexual selection. Here we examine the heritability and plasticity of a male dominance advertisement in the bank vole, Clethrionomys glareolus, in stable and…

Initial evolution of warning coloration: comments on the novel world method.

BLOOD PARASITES AND MALE FITNESS IN THE PIED FLYCATCHER

In vertebrates the effect of parasites on host ecology has almost been ignored. Recently the view that well-adapted parasites do not harm their hosts has been challenged and there is growing evidence that parasites do have a present-day effect on a great variety of host fitness components. The pied flycatcher is a small migratory passcrine bird. Any decrease in condition caused by disease should affect its ability to cope with physical demands of migration. Here we examine whether blood parasites have any effect on male arrival time. Males infected with Trypanosoma arrived on average 2 days later than males with no Trypanosoma infection. Infected males also had shorted tails and tended to h…

Polygyny and Breeding Success of Pied Flycatchers Nesting in Natural Cavities

Many hole-nesting passerine birds accept, or even prefer, nest boxes. Hence most of our knowledge about the breeding biology, population dynamics, life-history evolution and so forth of such species comes from nest box studies. This, in some cases, might lead to erroneous conclusions. For example, breeding density in nest box areas may often be much higher than in the natural situation (e.g. von Haartman, 1971), possibly leading to unnatural density dependent effects. Furthermore, the use of nest boxes may reduce predation risks (e.g. Nilsson 1975, 1984 a,b, Moller 1989), while the routine procedure of cleaning boxes after each breeding season may reduce the number of parasites, thus affect…

Viability costs of condition-dependent sexual male display in a drumming wolf spider

According to the conditional handicap models females use male ornaments as honest signals of male viability. The assumptions for honest signalling are that the traits are costly and that they reflect male phenotypic condition, and hence optimal trait size is largest in the most viable males. However, experimental evidence for the costs of signalling are scarce. In this study we experimentally tested whether acoustic signalling, drumming, in a wolf spider Hygrolycosa rubrofasciata is a condition dependent, costly trait, and thus offers an honest signal of quality to females. Males of this species court females by drumming dry leaves with their abdomen. Females prefer to mate with males of hi…

Sexual signalling and viability in a wolf spider ( Hygrolycosa rubrofasciata ): measurements under laboratory and field conditions

This study examined the crucial prediction of the conditional-handicap theory, the relationship between male sexual trait size and male viability, in the wolf spider Hygrolycosa rubrofasciata. In this species, males court females by drumming dry leaves with their abdomen, and males with the highest drumming rate enjoy highest mating success. We determined male drumming rate, body mass, and mobility, which reflects mate-searching activity, in relation to male survival. Because it is often difficult to know how results obtained from laboratory studies reflect the natural world, particularly when the measured variable is survival, we repeated our study in both laboratory and field conditions. …

Male mating success and risk of predation in a wolf spider: a balance between sexual and natural selection?

1.  Traits that benefit males through sexual selection are simultaneously expected to impair males by provoking costs through natural selection. If we consider the two male fitness components, mating success and viability, then we may expect that the increase in male mating success resulting from a larger trait size will be counterbalanced by an increase in viability costs. 2.  We studied the benefits and costs of male mate searching and sexual signalling activity in the wolf spider Hygrolycosa rubrofasciata. In the field, males search females actively and court them by drumming dry leaves with their abdomen. Females have been shown to prefer males with high drumming rate. Male moving and e…

Adaptive significance of synchronous chorusing in an acoustically signalling wolf spider

Synchronous sexual signalling is a behavioural phenomenon that has received considerable theoretical interest, but surprisingly few empirical tests have been conducted. Here, we present a set of experiments designed to determine (i) whether the sexual signalling of the drumming wolf spider Hygrolycosa rubrofasciata is synchronous, and (ii) whether the synchrony may have evolved through female preference. Using controlled playback experiments, we found that males actively synchronized their drumming bouts with other males and females significantly preferred closely synchronized drumming clusters compared with loose clusters. In loose clusters, the first drumming signals attracted the most fe…

Lekking in the black grouse— a test of male viability

LEKS, where males congregate to display and females attend only to mate, present one of the most remarkable outcomes of sexual selection1. It is a common but untested belief that females mate with the most vigorous males2. In leks of the black grouse Tetrao tetrix, males dominant in winter flocks were most successful in mating, as were males winning fights over female dummies placed at territory boundaries. Males tear feathers from each others' tail ornaments in combats, and attractive males always had undamaged tails. We report here that by choosing victorious males, females mate with males that are most likely to survive the following six months. There is a strong association between fema…

Maternal effort and male quality in the bank vole, Clethrionomys glareolus.

Parental investment in reproduction is adjusted according to potential benefits in terms of offspring survival and/or mating success. If male quality affects the reproductive success of a female, then females mating with high-quality males should invest more in reproduction. Although the subject has been of general interest, further experimental verification of the hypothesis is needed. We studied whether female bank voles (Clethrionomys glareolus) adjusted their maternal effort according to male quality, measured as mating success. To enable the measurement of maternal effort during nursing separately from male genetic effects the litters were cross-fostered. Further, the genetic backgroun…

The role of females in influencing mating patterns

Maternal antibodies in a wild altricial bird: effects on offspring immunity, growth and survival

1. In many animals immunity is not fully developed until adulthood but the young still need protection against various sets of pathogens. Thus, bird nestlings are highly dependent on antibodies received from their mother (in the eggs) during their rapid early growth period. The relationship between maternal immunity and the development of neonates' own immunity has been poorly studied. 2. It has been suggested that immune function plays an important part in mediating resource competition between different life-history traits, e.g. growth and reproduction. Maternal investment of antibodies has potentially permanent effects on offspring phenotype. Thus, the trade-offs between the immune funct…

Honesty of agonistic signalling and effects of size and motivation asymmetry in contests

Game theoretical models predict that the main function of fighting behaviour is to assess the relative fighting ability of opponents. The sequential assessment game has often been used to investigate contests, while honest signalling theory has received much less attention. With the wolf spider Hygrolycosa rubrofasciata we investigated whether male agonistic signalling can reveal honest information about fighting ability, and how size and motivation asymmetries affect male fighting behaviour. We also determined whether male–male competition affects the courtship behaviour of the males. We found that agonistic drumming activity is an honest indicator of male fighting ability, and that relati…

Female-female aggression and female mate choice on black grouse leks.

We studied female-female aggression in relation to female mate choice in black grouse, Tetrao tetrix, in central Finland, in 1994-1998. Aggression occurred on average every other minute when there was more than one female on a territory, and aggressive behaviour was most prominent when several females attended the lek. Interactions tended to be proportionally most frequent on the territories of the highest-ranking males, although not significantly so. Females that were chased by other females did not mate with lower-ranking males than their aggressors did. Furthermore, chased females were only rarely (6% of cases) forced to move off the territory by agonistic interactions and copulations we…

Trade-offs between sexual advertisement and immune function in the pied flycatcher (Ficedula hypoleuca).

Good genes models of sexual selection assume that sexual advertisement is costly and thus the level of advertisement honestly reveals heritable viability. Recently it has been suggested that an important cost of sexual advertisement might be impairment of the functioning of the immune system. In this field experiment we investigated the possible trade-offs between immune function and sexual advertisement by manipulating both mating effort and activity of immune defence in male pied flycatchers. Mating effort was increased in a non-arbitrary manner by removing females from mated males during nest building. Widowed males sustained higher haematocrit levels than control males and showed higher…

Fluctuating asymmetry and copulation success in lekking black grouse

Abstract In lekking black grouse, Tetrao tetrix males at the centre of the leks obtain more copulations than males at the edges. We found that males with territories at the edge of the lek obtained fewer matings and also have the most asymmetric tarsi. However, when considering the tail ornament (the lyre) no correlation between asymmetry and mating success was found. Although females are unlikely to select males on the basis of tarsus symmetry, asymmetry in this trait may reflect male condition. We suggest that males in poor condition, as reflected by higher levels of fluctuating asymmetry in the tarsi, do not achieve central positions on the lek arena. This would account for the lower suc…

Predator experience on cryptic prey affects the survival of conspicuous aposematic prey.

Initially, aposematism, which is an unprofitable trait, e.g. noxiousness conspicuously advertised to predators, appears to be a paradox since conspicuousness should increase predation by naive predators. However, reluctance of predators for eating novel prey (e.g. neophobia) might balance the initial predation caused by inexperienced predators. We tested the novelty effects on initial predation and avoidance learning in two separate conspicuousness levels of aposematic prey by using a 'novel world' method. Half of the wild great tits (Parus major) were trained to eat cryptic prey prior to the introduction of an aposematic prey, which potentially creates a bias against the aposematic morph. …

Decreased sexual signalling reveals reduced viability in small populations of the drumming wolf spider Hygrolycosa rubrofasciata.

One of the important goals in conservation biology is to determine reliable indicators of population viability. Sexual traits have been suggested to indicate population extinction risk, because they may be related to viability through condition dependence. Moreover, condition-dependent sexual traits may be more sensitive indicators of population viability than early life-history traits, because deleterious fitness effects of inbreeding tend to be expressed mainly at the end of the species' life history. However, empirical evidence of the significance of sexual behaviour for population viability is missing. In this study, we examined two male sexual traits and survival in 39 different-sized …

Individual and Environmental Determinants of Daily Black Grouse Nest Survival Rates at Variable Predator Densities

Nest predation in ground nesting black grouse (Tetrao tetrix) inhabiting managed boreal forests is arguably the single most important cause of nesting failure. Little is known, however, about indirect effects of other factors, such as maternal or environmental properties, and to what extent maternal and habitat qualities interact with varying levels of predator densities. Using an information-theoretical approach, we studied maternal and environmental determinants of daily nest survival rates under variable predator densities of 210 individual black grouse hens in central Finland. Environmental determinants were far more important than maternal ones, and the effects were more apparent at hi…

Female-biased dispersal alone can reduce the occurrence of inbreeding in black grouse (Tetrao tetrix)

Although inbreeding depression and mechanisms for kin recognition have been described in natural bird populations, inbreeding avoidance through mate choice has rarely been reported suggesting that sex-biased dispersal is the main mechanism reducing the risks of inbreeding. However, a full understanding of the effect of dispersal on the occurrence of inbred matings requires estimating the inbreeding risks prior to dispersal. Combining pairwise relatedness measures and kinship assignments, we investigated in black grouse whether the observed occurrence of inbred matings was explained by active kin discrimination or by female-biased dispersal. In this large continuous population, copulations b…

Tracking the evolution of warning signals

EVOLUTIONARYstudies are hampered by a lack of experimental ways in which to test past events such as the origination of aposematism1–7, whereby unpalatable or poisonous prey signal their unprofitability, often by being warningly coloured. Inexperienced predators do learn to avoid unpalatable prey as a result of such signals8–10, but in addition there may be an inherited cautiousness about attacking when common or conspicuous warning signals are evident11–16. As current predators are not naive in the evolutionary sense, it is still not resolved3–7,17,18 whether aposematism originated only in aggregations of prey19,20 or among solitary prey as well21–23. Here we explore this controversy in ev…

Batesian Mimicry and Signal Accuracy

RICE, W. R. 1989. Analyzing tables of statistical tests. Evolution 43:223-225. RICE, W. R., AND E. E. HOSTERT. 1993. Laboratory experiments on speciation: what have we learned in 40 years? Evolution 47: 1637-1653. SAWADA, S. 1963. Studies on the local races of the Japanese newt, Triturus pyrrhogaster Boie. II. Sexual isolation mechanisms. J. Sci. Hiroshima Univ. Ser. B 21:167-180. SPIETH, H. T, AND J. M. RINGO. 1983. Mating behavior and sexual isolation in Drosophila. Pp 223-284 in M. Ashburner, L. M. Carson, and J. N. Thompson Jr, eds. The genetics and biology of Drosophila. Academic Press, New York. TEMPLETON, A. R. 1996. Experimental evidence for the genetictransilience model of speciati…

Mate choice for offspring performance: major benefits or minor costs?

There is considerable disagreement over whether or not gaining viability benefits to offspring could be substantial enough to overcome the costs of female choosiness. A recent review suggested that the 'lek paradox' might be resolved by large indirect benefits as indicated by highly heritable ornamental traits. We selected males of a wolf spider Hygrolycosa rubrofasciata in relation to their sexual signalling rate (audible drumming). The estimated correlated response in offspring viability was rather small (0.12 s.d.). However, it may be large enough if the costs of being choosy are small. In fact, females mate with better-than-average males just by responding passively to a random drumming…

Black Grouse leks on ice: Female mate sampling by incitation of male competition?

Male-male competition is assumed to limit female choice of mates, but it may also help females to choose the most vigorous males. We studied the mate sampling behaviour of female black grouse (Tetrao tetrix) at spatially unstable leks on ice-covered lakes. In the absence of territories and site-dependence in outcomes of fights, the male dominance hierarchy is very evident on ice. When being courted by dominant males, females frequently tried to approach other males. This was frequently prevented because (1) the courting male and the approached male were involved in physical fight, or (2) the dominant male followed the female and the approached male escaped and avoided contact with him. Thes…

THE EFFECT OF ALTERNATIVE PREY ON THE DYNAMICS OF IMPERFECT BATESIAN AND MÜLLERIAN MIMICRIES

Both Batesian and Müllerian mimicries are considered classical evidence of natural selection where predation pressure has, at times, created a striking similarity between unrelated prey species. Batesian mimicry, in which palatable mimics resemble unpalatable aposematic species, is parasitic and only beneficial to the mimics. By contrast, in classical Müllerian mimicry the cost of predators' avoidance learning is shared between similar unpalatable co-mimics, and therefore mimicry benefits all parties. Recent studies using mathematical modeling have questioned the dynamics of Müllerian mimicry, suggesting that fitness benefits should be calculated in a way similar to Batesian mimicry; that i…

Combs and sexual selection in black grouse (Tetrao tetrix)

We studied supra-orbital combs in lekking black grouse (Telrao tetrix) in relation to sexual selection at five leks in Finland 1991-1998 and four leks in Sweden 1992-1995. Comb size was estimated in two ways: by observing its natural size in the field at

Negatively condition dependent predation cost of a positively condition dependent sexual signalling.

Predation is considered as an important factor constraining the expression of sexual signals. Nevertheless, direct quantitative evidence for predation provoking significant viability costs on individuals signalling at high rates is scarce. Moreover, it is unclear whether high rate signallers are able to balance presumably increased predation costs. We examined whether a condition dependent audible sexual signal, drumming, makes Hygrolycosa rubrofasciata male spiders more prone to predation by pied flycatchers (Ficedula hypoleuca), and whether sexual signalling rate is related to escaping ability once attacked. When birds were given a choice between two spider males manipulated to drum eithe…

Patterns of variation in tail ornament size in birds

In recent years several different kinds of sexual selection models have been developed, and tail ornaments in birds have frequently been used as an example of a sexually selected character where the models might apply. However, very little is known about intra- and interpopulation variation in ornament size. We have studied the elongated tail ornaments in four species of whydahs Vidua, the forktailed flycatcher Tyrannus savana and the Asian paradise flycatcher Terpsiphone paradisi. Ornaments were relatively longer in males with the longest tarsi (‘heterogony’ with positive allometry). Also, tail lengths were remarkably variable within each geographical area, the coefficient of variation (av…

Color Signals in the Black Grouse (Tetrao tetrix): Signal Properties and Their Condition Dependency

Recent progress in techniques of quantifying between-individual differences of color-based ornaments has revealed undiscovered possibilities for research in sexual selection. We present how the color spectra data can be comprehensively used for studying the importance of sexual ornaments in the black grouse and how these ornaments are related to a male condition. For this, we used both correlative field and experimental data. Field data indicated that older males had more chromatic coloration than yearlings. Blue chroma of males was correlated with male mating success. We experimentally manipulated yearling birds with testosterone implants and found that testosterone-implanted males had imp…

Data from: Limited indirect fitness benefits of male group membership in a lekking species

In group living species, individuals may gain the indirect fitness benefits characterising kin selection when groups contain close relatives. However, tests of kin selection have primarily focused on cooperatively breeding and eusocial species, whereas its importance in other forms of group living remains to be fully understood. Lekking is a form of grouping where males display on small aggregated territories, which females then visit to mate. As females prefer larger aggregations, territorial males might gain indirect fitness benefits if their presence increases the fitness of close relatives. Previous studies have tested specific predictions of kin selection models by using measures such …

Data from: Life history differences in age-dependent expressions of multiple ornaments and behaviors in a lekking bird

Age is a major factor explaining variation in life-history traits among individuals with typical patterns of increasing trait values early in life, maximum trait expression, and senescence. However, age-dependent variation in the expressions of sexually selected traits has received less attention, although such variation underpins differences in male competitive abilities and female preference, which are central to sexual selection. In contrast to previous studies focusing on single traits, we used repeated measures of seven sexually selected morphological and behavioral traits in male black grouse (Tetrao tetrix) to quantify the effects of age and life span on their expressions and quantif…