Indicators of physiological stress and the elaboration of sexual traits in the collared flycatcher

Stress may have consequences for the evolution of condition-dependentsexual traits. For example, stress may be related to sexualtraits through immune function, and sexual traits can reflecthow individuals bear the costs of stress-mediated immunosuppression.However, male traits may be directly associated with stress,and such traits would then indicate stress tolerance. Here,we present initial results for the relationship between physiologicalstress estimated by the levels of heat shock proteins (HSP60and HSP70) and heterophil/lymphocyte ratio and the elaborationof sexual traits, such as forehead and wing patch size and songfeatures in the collared flycatcher Ficedula albicollis. Malesproducing longer and more versatile songs had significantlyhigher levels of HSP70, but other traits were unrelated to stress.In general, effect sizes for the relationship between stressand sexual traits had broad confidence intervals and variedbetween being small and medium effects. Immunoglobulin levels,leukocyte abundance, haemoparasite prevalence, male age, anddate and time effects did not affect the relationship betweenstress and sexual traits. These preliminary results, servinga basis for further experimental studies indicate that the relationshipbetween sexual traits and stress does not seem to be strong,but stress may partially constrain the expression of some sexualtraits.     

Elevated spring testosterone increases parasite intensity in male red grouse

The expression of testosterone-dependent sexual traits mightsignal the ability of their bearers to cope with parasite infections.According to the immunocompetence handicap hypothesis (IHH),such signals would be honest because physiological costs oftestosterone, such as a reduced ability to control parasiteinfections, would prevent cheating. We tested whether testosteronewould affect the outcome of a standardized parasite challengein red grouse, using a main parasite of the species, the nematodeTrichostrongylus tenuis. We caught males in spring, removedtheir nematode parasites, and implanted them with testosteroneor empty implants, as controls. After 1 month, they were reinfectedwith a standard dose of infective T. tenuis parasites. Whenchallenged, testosterone males had relatively less globulinrelative to albumin plasma proteins than control males, an indicationthat they had experienced increased physiological stress. Testosterone-treatedmales had significantly more T. tenuis parasites than controlsin the next autumn and also had more coccidia and lost moreweight than controls. Testosterone-treated males neverthelessbenefited from their elevated spring testosterone: they hadbigger sexual ornaments than controls both in spring and autumn,and they tended to have a higher pairing and breeding successthan controls. Our results supported the IHH in showing thatelevated testosterone impaired the ability of males to copewith a standardized challenge by a dominant parasite. Testosteronethus plays a key role in mediating trade-offs between reproductiveactivities and parasite defense, and testosterone-dependentcomb size might honestly signal the ability of red grouse tocontrol T. tenuis infection.     

The evolution of immune defense and song complexity in birds

Abstract There are three main hypotheses that explain how the evolution of parasite virulence could be linked to the evolution of secondary sexual traits, such as bird song. First, as Hamilton and Zuk proposed a role for parasites in sexual selection, female preference for healthy males in heavily parasitized species may result in extravagant trait expression. Second, a reverse causal mechanism may act, if sexual selection affects the coevolutionary dynamics of host-parasite interactions per se by selecting for increased virulence. Third, the immuno-suppressive effects of ornamentation by testosterone or limited resources may lead to increased susceptibility to parasites in species with elaborate songs. Assuming a coevolutionary relationship between parasite virulence and host investment in immune defense we used measures of immune function and song complexity to test these hypotheses in a comparative study of passerine birds. Under the first two hypotheses we predicted avian song complexity to be positively related to immune defense among species, whereas this relationship was expected to be negative if immuno-suppression was at work. We found that adult T-cell mediated immune response and the relative size of the bursa of Fabricius were independently positively correlated with a measure of song complexity, even when potentially confounding variables were held constant. Nestling T-cell response was not related to song complexity, probably reflecting age-dependent selective pressures on host immune defense. Our results are consistent with the hypotheses that predict a positive relationship between song complexity and immune function, thus indicating a role for parasites in sexual selection. Different components of the immune system may have been independently involved in this process.     

A partly coverable badge signalling avian virus resistance

We investigated whether the sexually selected forehead patch of the collared flycatcher Ficedula albicollis is an honest badge of status indicating quality expressed as immunological response. We used both manual measurements and digital measurements, the latter based on photographs. Badge‐size data were collected during the mating period and during the nestling feeding period to capture trait plasticity. Concomitant with first sample collection, birds were inoculated with a novel antigen. Antibody response was strongly and positively correlated with badge expression during the mating period and with the increase in badge expression during the mating period as compared with outside this period. The results support the Hansen and Rohwer theory of coverable badges, are consistent with the immunocompetence handicap hypothesis and with the good genes model suggesting that, on a population level, the expression of secondary sexual traits should be an honest signal positively associated with traits that are beneficial for survival. The results also suggest that manual measurements of this type of secondary sexual trait are sufficiently exact.     

POPULATION DIVERGENCE IN SEXUAL ORNAMENTS: THE WHITE FOREHEAD PATCH OF NORWEGIAN PIED FLYCATCHERS IS SMALL AND UNSEXY

Models of sexual selection suggest that populations may easily diverge in male secondary sexual characters. Studies of a Spanish population of the pied flycatcher, Ficedula hypoleuca, and a Swedish population of the closely related collared flycatcher, F. albicollis, have indicated that the white forehead patch of males is a sexually selected trait. We studied the white forehead patch of male pied flycatchers (n = 487) in a Norwegian population over seven years. Males with large forehead patches were in general more brightly colored, but patch height was not correlated to body mass, body size, or parasite loads. Conditions during the nestling period did not seem to influence patch height as an adult. Patch height increased slightly from the first to the second year as adults, but then remained relatively constant at higher ages. Patch height was not related to survival. Year-to-year changes showed that males who increased in patch height also increased in body mass, suggesting that expression of the forehead patch may be partly condition dependent. However, changes in body mass explained only a small proportion of the variance in patch height between males. Thus, patch height would not be a good indicator of male quality. Furthermore, patch size was also not related to male ability to feed nestlings, indicating that females would not obtain direct benefits by choosing males with large patches. However, patch height could be a Fisher trait, but this requires heritability and there was no significant father-son resemblance in patch height. Comparisons of the males visited by each female during the mate sampling period indicated that chosen males did not have larger forehead patches than rejected males. Experimental manipulation of patch height did not affect male mating success. These results indicate that females do not use patch size as a mate choice cue. Finally, patch height did not predict the outcome of male contests for nestboxes, suggesting that the forehead patch is not an intrasexually selected cue of status. Norwegian pied flycatchers have smaller forehead patches than both Spanish pied flycatchers and Swedish collared flycatchers. We suggest that this pattern may be explained by the lack of sexual selection on the forehead patch in the Norwegian population as compared to the other populations, where the patch is apparently sexually selected. We discuss possible reasons for these population divergences, such as female choice on an alternative secondary sexual character (general plumage color) and speciation among Ficedula flycatchers.     

Hamilton and Zuk meet heterozygosity? Song repertoire size indicates inbreeding and immunity in song sparrows (Melospiza melodia)

Hamilton and Zuk's influential hypothesis of parasite-mediated sexual selection proposes that exaggerated secondary sexual ornaments indicate a male's addictive genetic immunity to parasites. However, genetic correlated of ornaments and immunity have rarely been explicitly identified. Evidence supporting Hamilton and Zuk's hypothesis has instead been gathered by looking for positive phenotypic correlations between ornamentation and immunity; such correlations are assumed to reflect causal, addictive relationships between these traits. We show that in a song sparrows, Melospiza melodia, male's song repertoire size, a secondary sexual trait, increased with his cell-mediated immune response (CMI) to an experimental challenge. However, this phenotypic correlation could be explained because both repertoire size and CMI declined with a male's inbreeding level. Repertoire size therefore primarily indicated a male's relative heterozygosity, a non-addictive genetic predictor of immunity. Caution may therefore be required when interpreting phenotypic correlations as support for Hamilton and Zuk's addictive model of sexual selection. However, our results suggest that female song sparrows choosing with large repertoires would on average acquire more outbred and therefore more heterozygous mates. Such genetic dominance effects on ornamentation are likely to influence evolutionary trajectories of female choice, and should be explicitly incorporated into genetic models of sexual selection.     

Age-Dependent Expression of Song in the Collared Flycatcher, Ficedula albicollis

Theory suggests that male ornaments should be reliable signals of age, with more elaborated ornaments reflecting superior quality in terms of experience and/or viability. Bird song is immensely involved in sexual selection, thus not‐surprisingly, it usually shows age‐dependent variation. Although the collared flycatcher Ficedula albicollis has been intensively studied for its sexual traits, and female preference for male age is expected to be strong, there is no quantitative information on age‐dependent expression of song in this species. Here, we fill this gap and, based on phenotypic correlations, we report the relationship between age and several song features. Repertoire size was consistently smaller in yearlings than in older males, but it also tended to increase after the second year of breeding. In a meta‐analysis of effect sizes using data from the literature, we found that the strength of the relationship between age and repertoire size in the collared flycatcher is similar to patterns observed in other passerines. Song rate was inversely related to age, as yearlings sang at higher rates than older males. Generally, effect sizes for the relationship between age and other song traits were of medium level on average, and had broad confidence intervals. Song traits covaried with survival in a direction suggesting that differences in song between age categories are unlikely to result from phenotype dependent survival. Our results generally support the hypothesis that song holds the potential to function as a reliable indicator of male age in the collared flycatcher. However, it seems that not all song traits are unambiguously more expressed in older males than in yearlings.     

Immune system activation affects male sexual signal and reproductive potential in crickets

Parasite-mediated sexual selection theory posits that individuals(usually females) choose mates by assessing the expression ofcostly secondary sexual signals, which provide reliable indicationsof parasite resistance. If these signals are indeed reliable,then immune-compromised males are predicted to exhibit changesin the sexual signal that are discernable by the female. Moreover,the mating pair is predicted to exhibit some reduction in reproductivefitness if the male is immune compromised. Here, we addressedthese predictions in the ground cricket, Allonemobius socius,by injecting juvenile males with lipopolysaccarides, which allowedus to activate the immune system without the introduction ofa metabolically active pathogen. As a consequence, we were ableto disentangle the cost of immune system activation from thecost of infection. We found that immune activation had a long-termeffect on male calling song and the males' ability to providepaternal resources, which can constrain male and female reproductivepotential. We also found that song interpulse interval variedsignificantly with the male's immune treatment and may thereforeprovide choosy females with a way to avoid mating with immune-compromisedmales. In short, our data support the parasite-mediated theoryof sexual selection, suggesting that female's gain direct benefitsby mating with males who are immune competent.     

Genetic Architecture of Sexual Selection: QTL Mapping of Male Song and Female Receiver Traits in an Acoustic Moth

Models of indirect (genetic) benefits sexual selection predict linkage disequilibria between genes that influence male traits and female preferences, owing to non-random mate choice or physical linkage. Such linkage disequilibria can accelerate the evolution of traits and preferences to exaggerated levels. Both theory and recent empirical findings on species recognition suggest that such linkage disequilibria may result from physical linkage or pleiotropy, but very little work has addressed this possibility within the context of sexual selection. We studied the genetic architecture of sexually selected traits by analyzing signals and preferences in an acoustic moth, Achroia grisella, in which males attract females with a train of ultrasound pulses and females prefer loud songs and a fast pulse rhythm. Both male signal characters and female preferences are repeatable and heritable traits. Moreover, female choice is based largely on male song, while males do not appear to provide direct benefits at mating. Thus, some genetic correlation between song and preference traits is expected. We employed a standard crossing design between inbred lines and used AFLP markers to build a linkage map for this species and locate quantitative trait loci (QTL) that influence male song and female preference. Our analyses mostly revealed QTLs of moderate strength that influence various male signal and female receiver traits, but one QTL was found that exerts a major influence on the pulse-pair rate of male song, a critical trait in female attraction. However, we found no evidence of specific co-localization of QTLs influencing male signal and female receiver traits on the same linkage groups. This finding suggests that the sexual selection process would proceed at a modest rate in A. grisella and that evolution toward exaggerated character states may be tempered. We suggest that this equilibrium state may be more the norm than the exception among animal species.     

Song and immunological condition in male barn swallows (Hirundo rustica)

Male secondary sexual characters may have evolved as intra-or intersexual signals of male phenotypic or genetic quality.In birds, singing performance may have the function to honestlyreveal health and vigor of individual males. Infectious diseasesand poor body conditions would therefore be expected to negativelyinfluence singing performance. Since bird pathogens are knownto elicit both a humoral and a cell-mediated immune response,it can be predicted that a negative relationship exists betweensinging performance and activity of the immune system. Thisprediction was tested for the first time in this correlationalstudy. The relationships between song rate and features andhematological variables (concentration of leukocytes in peripheralblood, ratio of gamma-globulins to total plasma proteins, bloodcell sedimentation rate, hematocrit) and body condition wereanalyzed in a population of bam swallows (Hirundo rustica).Song rate was negatively correlated with lymphocyte concentrationand with the ratio of gamma-globulins to plasma proteins. Spectrographicanalysis showed that features of song were not significantlycorrelated with hematological variables or body condition. Thelevel of circulating testosterone was not correlated with songrate nor hematological variables. This study is the first toshow a correlation between a bird's singing performance andhematological profile and suggests that song rate of male barnswallows may reflect their health status. Song in this speciesmight thus have evolved because it allows prospecting femalesto assess aspects of phenotypic and/or genetic quality of potentialmates     

Oxidative damage and plasma antioxidant capacity in relation to body size,age, male sexual traits and female reproductive performance in the collared flycatcher (<Emphasis Type="Italic">Ficedula albicollis</Emphasis>)

The study of oxidative stress is a potential tool for studying the functional interactions among life history traits, sexual traits and physiological status in animals. In this study, we investigated relationships between measures of plasma oxidative status and male sexual traits, female reproductive investment and three other life history traits, in a wild population of collared flycatchers (Ficedula albicollis). Flycatcher males with a larger white forehead patch had higher level of plasma antioxidant capacity. For females, clutch size was not associated with plasma oxidative status, but egg size was positively correlated with antioxidant capacity. The relationship between age and levels of plasma oxidative damage remains controversial in this species: young female flycatchers showed higher levels of hydroperoxides compared to antioxidants, whereas age did not predict oxidative status of males. Males had higher levels of oxidative damage than females, although the concentration of antioxidant compounds was similar between the sexes. Females that mated with more ornamented males had higher plasma antioxidant capacity. Our results suggest that, for males and females, greater investment in sexual signal and reproduction, respectively, does not reduce the capacity for self-maintenance or avoidance of oxidative stress. Finally, our data support indirectly the occurrence of assortative mating in our species, since females with higher plasma antioxidant capacity mated with more ornamented males.     

Acoustic preference functions and song variability in the Hawaiian cricket Laupala cerasina

Female preference functions for different sexual traits can differ significantly, from 'unimodal' to 'open ended'. Through the study of acoustic communication in anurans, several studies have reported an association between static (stereotyped) traits versus dynamic (variable) traits and preference function shape (unimodal versus open ended, respectively). Observing a similar pattern in a phylogenetically independent group would suggest that deterministic forces have caused a relationship between signal variability and preference function shape in acoustic signalling systems. We examined this phenomenon in crickets, another animal characterized by intersexual acoustic communication. We measured the within-male variability for three acoustic features of the male calling song in Laupala cerasina and the corresponding shape of the female preference function for each of these features. We offer support for the generalization that open-ended preference functions correspond to relatively dynamic courtship traits and unimodal preference functions correspond to relatively static courtship traits. We discuss the evolutionary significance of these findings in the context of the natural history of the Laupala species radiation.     

Environmental variability and acoustic signals: a multi-level approach in songbirds

Among songbirds, growing evidence suggests that acoustic adaptation of song traits occurs in response to habitat features. Despite extensive study, most research supporting acoustic adaptation has only considered acoustic traits averaged for species or populations, overlooking intraindividual variation of song traits, which may facilitate effective communication in heterogeneous and variable environments. Fewer studies have explicitly incorporated sexual selection, which, if strong, may favour variation across environments. Here, we evaluate the prevalence of acoustic adaptation among 44 species of songbirds by determining how environmental variability and sexual selection intensity are associated with song variability (intraindividual and intraspecific) and short-term song complexity. We show that variability in precipitation can explain short-term song complexity among taxonomically diverse songbirds, and that precipitation seasonality and the intensity of sexual selection are related to intraindividual song variation. Our results link song complexity to environmental variability, something previously found for mockingbirds (Family Mimidae). Perhaps more importantly, our results illustrate that individual variation in song traits may be shaped by both environmental variability and strength of sexual selection.     

The blue tit's song is an inconsistent signal of male condition

Sexually selected traits are often hypothesized to signal malecondition or quality, though empirical evidence is mixed, anda number of alternative models of sexual selection do not requirecondition dependence. We examined the relationship between variousmeasures of condition and dawn songs in male blue tits (Cyanistescaeruleus). We detected 6 largely independent measures of variation(i.e., variables) in these songs. None of these variables wererelated to blue tits' ultraviolet–blue plumage, a demonstratedsexual signal, thus failing to support the redundant signalhypothesis. We found some evidence that the song variables wemeasured signaled male quality. There were correlations betweenbody size and certain song traits, though neither male age normale recapture in the subsequent breeding season (apparent localsurvival) predicted any song variation. We combined our resultswith published effect sizes comparing blue tit song with malequality variables using meta-analysis and found that a few songmeasures are correlates of male quality, though as in our fielddata, neither male age nor survival appeared related to song.Our relatively large sample sizes (>60), combined with ourmeta-analytical integration of 89 effect sizes, make the resultsregarding the signaling value of our measured components ofblue tit song robust. These results demonstrate that 1) onlycertain aspects of signal variation may be condition dependentand 2) even when components of a sexual signal appear correlatedwith condition in some studies, these signal components maybe unrelated or inconsistently related to a variety of conditionindices.     

Flycatcher song in allopatry and sympatry--convergence, divergence and reinforcement

The theory of reinforcement predicts that natural selection against the production of unfit hybrids favours traits that increase assortative mating. Whether culturally inherited traits, such as bird song, can increase assortative mating by reinforcement is largely unknown. We compared songs of pied (Ficedula hypoleuca) and collared flycatchers (F. albicollis) from two hybrid zones of different ages with songs from allopatric populations. Previously, a character divergence in male plumage traits has been shown to reinforce premating isolation in sympatric flycatchers. In contrast, we find that the song of the pied flycatcher has converged towards that of the collared flycatcher (mixed singing). However, a corresponding divergence in the collared flycatcher shows that the species differences in song characters are maintained in sympatry. Genetic analyses suggest that mixed song is not caused by introgression from the collared flycatcher, but rather due to heterospecific copying. Circumstantial evidence suggests that mixed song may increase the rate of maladaptive hybridization. In the oldest hybrid zone where reinforcement on plumage traits is most pronounced, the frequency of mixed singing and hybridization is also lowest. Thus, we suggest that reinforcement has reduced the frequency of mixed singing in the pied flycatcher and caused a divergence in the song of the collared flycatcher. Whether a culturally inherited trait promotes or opposes speciation in sympatry may depend on its plasticity. The degree of plasticity may be genetically determined and accordingly under selection by reinforcement.     

Song post exposure, song features, and predation risk

Male birds use song to attract mates and deter other males,but in doing so, they also attract the attention of predatorsand parasites. Such viability costs are inherent in reliablesignals, potentially causing females to prefer mates that displayfrom the most exposed sites. However, viability costs of sexualsignals may be ameliorated by affecting the choice of microhabitat,which in turn may affect the design of song features that aremost efficiently transmitted in this microhabitat. We estimatedthe exposure of song posts (microsites used by males when singing)used by passerine birds in relation to prey selection by thesparrowhawk Accipiter nisus, by calculating the proportion ofmales that sang from song posts that were at the maximum levelof the vegetation, in an attempt to quantify the costs of sexualselection. We quantified prey susceptibility to predation asthe difference between the log-transformed observed number ofprey minus the log-transformed expected number of prey in theenvironment. This prey susceptibility index increased with increasingsong post exposure similarly in sexually dichromatic and monochromaticspecies, although the prey susceptibility index was relatedto sexual dichromatism. Song post exposure was dependent onhabitat, but comparative models controlling for the potentiallyconfounding effects of habitat, sexual dichromatism, hole nesting,coloniality, body mass, cognitive capacities, and flying abilitiesindicated that the relationship between the prey susceptibilityindex and song post exposure is strong. Path analyses of therelationship between song post exposure, sexual dichromatism,and prey susceptibility index revealed that selection actingon sexual dichromatism and song post exposure has secondaryimpact on prey susceptibility index. The opposite causal mechanismsby which predation affects sexual traits are less likely. Thesemodels suggest that female preference for high song posts ordichromatic plumage increases predation risk on an evolutionarytime scale.     

Climatic and temporal effects on the expression of secondary sexual characters: genetic and environmental components

Despite great interest in sexual selection, relatively little is known in detail about the genetic and environmental determinants of secondary sexual characters in natural populations. Such information is important for determining the way in which populations may respond to sexual selection. We report analyses of genetic and large-scale environmental components of phenotypic variation of two secondary sexual plumage characters (forehead and wing patch size) in the collared flycatcher Ficedula albicollis over a 22-year period. We found significant heritability for both characters but little genetic covariance between the two. We found a positive association between forehead patch size and a large-scale climatic index, the North Atlantic Oscillation (NAO) index, but not for wing patch. This pattern was observed in both cross-sectional and longitudinal data suggesting that the population response to NAO index can be explained as the result of phenotypic plasticity. Heritability of forehead patch size for old males, calculated under favorable conditions (NAO index > or = median), was greater than that under unfavorable conditions (NAO index < median). These changes occurred because there were opposing changes in additive genetic variance (VA) and residual variance (VR) under favorable and unfavorable conditions, with VA increasing and VR decreasing in good environments. However, no such effect was detected for young birds, or for wing patch size in either age class. In addition to these environmental effects on both phenotypic and genetic variances, we found evidence for a significant decrease of forehead patch size over time in older birds. This change appears to be caused by a change in the sign of viability selection on forehead patch size, which is associated with a decline in the breeding value of multiple breeders. Our data thus reveal complex patterns of environmental influence on the expression of secondary sexual characters, which may have important implications for understanding selection and evolution of these characters.     

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 expression of forehead patch height, suggesting that manipulation succeeded in increasing mating effort. Males that were experimentally forced to increase mating effort had reduced humoral immune responsiveness compared with control males. In addition, experimental activation of immune defence by vaccination with novel antigens reduced the expression of male ornament dimensions. To conclude, our results indicate that causality behind the trade-off between immune function and sexual advertisement may work in both directions: sexual activity suppresses immune function but immune challenge also reduces sexual advertisement.     

Rapid temporal change in the expression and age-related information content of a sexually selected trait

The expression of sexual signals is often phenotypically plastic and also evolves rapidly. Few studies have considered the possibility that proximate determination -- the pathway between genes and trait expression -- may also be subject to both phenotypic plasticity and evolutionary change. We examined long-term patterns in size, condition- and age-dependence, repeatability and heritability of forehead patch size, a sexually selected plumage trait in male collared flycatchers. We also estimated survival and sexual selection on the phenotypic value of the trait. Forehead patch size linearly declined during the 15 years, probably due to the significantly negative survival selection. In addition, the expression of genetic variation for the ornament apparently underwent an age-limited change, which implies a change in the information content of the signal to receivers. The persistent lack of condition-dependence makes phenotypic plasticity an unlikely explanation to our results. This raises the possibility of a microevolutionary change of both expression and proximate determination during the study period.     

Testosterone, immunocompetence, and honest sexual signaling in male red grouse

The expression of sexual ornaments has been suggested to reliablyindicate individual quality, such as the ability to cope withparasites and diseases. The Immunocompetence Handicap Hypothesis(IHH) states that testosterone-dependent ornaments honestlysignal such quality because of physiological costs associatedwith testosterone, such as impaired immune function. We testedpredictions of the IHH both correlatively and experimentallyin red grouse Lagopus lagopus scoticus. Male grouse exhibitsupra-orbital red combs whose size is testosterone-dependent.We found that comb size was not correlated to infection intensityby two parasites (coccidia and the nematode Trichostrongylustenuis), but it was significantly positively correlated withcondition and T-cell-mediated immunity (the ability to mounta primary inflammatory response). We manipulated testosteroneby means of implants and re-caught males after a month to investigatethe effects on comb size, condition, immunity, and parasiteload. Males implanted with testosterone had increased comb size,lost more condition, and had lower T-cell-mediated immunitythan control males. Increased testosterone also resulted ina significant increase in coccidia infection intensity but hadno effect on T. tenuis burden. The results are consistent withpredictions of the IHH and suggest that comb size honestly indicatesimmunocompetence and males' ability to cope with certain parasites.Females could thus benefit from choosing mates based on theexpression of this sexual trait.