Friendships between males and lactating females in a free-ranging group of olive baboons (Papio hamadryas anubis): evidence from playback experiments

Close association between an anoestrous female at the time of lactation and adult male(s) is relatively rare in mammals, but common in baboons (Papio hamadryas subsp.). The functional significance of these “friendships” remains unclear, however. In chacma baboons (P. h. griseipes), friendships are a counter-strategy to infanticide by immigrant males. Experimental playback of female distress calls in chacma baboons revealed that male friends are more motivated to protect females and infants than are control males. Olive baboons (P. h. anubis) also exhibit friendships, but infanticide is rare, suggesting that friendships provide females with protection from non-lethal aggression (anti-harassment hypothesis) or serve to promote male–infant bonds that later benefit the maturing juvenile (future male caretaker hypothesis). We replicated these playback experiments on a group of olive baboons to test between these hypotheses and to evaluate if the lower costs of non-lethal harassment lessens male protective responsiveness relative to protection from (more costly) infanticide. Spatial data revealed that most lactating females had one to four friend males. Relative to non-friends, friend dyads were characterized by higher rates of allogrooming and infant handling, but less agonism. Female rank was correlated with the number of male friends. Just as in chacma baboons, playback of female screams elicited stronger responses from male friends than control males in support the anti-harassment hypothesis. Compared to the chacma baboon, male olive baboons appeared to exhibit similarly high levels of protective solicitude for female friends although they protect against non-lethal harassment rather than infanticide.     

Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, <Emphasis Type="Italic">Scaphiopus couchii</Emphasis>

Females across many taxa commonly use multiple or complex traits to choose mates. However, the functional significance of multiple or complex signals remains controversial and largely unknown. Different elements of multiple or complex signals may convey independent pieces of information about different aspects of a prospective mate (the “multiple messages” hypothesis). Alternatively, multiple or complex signals could provide redundant information about the same aspect of a prospective mate (the “redundant” or “back-up” signal hypothesis). We investigated these alternatives using spadefoot toads, Scaphiopus couchii. Spadefoot toads primarily use calls to attract their mates, but males also exhibit sexually dimorphic coloration. We investigated whether male coloration is indicative of male size, condition, or infection status by a socially transmitted monogenean flatworm. We found that male coloration and dorsal patterning predicts male size and condition but not infection status. Moreover, when we presented females with a choice between a bright male model and a dark male model, we found that females preferred the bright model. Because aspects of males’ calls are also associated with male size and condition, we conclude that coloration is a potentially redundant indicator of male phenotype. We suggest that coloration could enhance mate choice in conjunction with male calling behavior by providing females with a long distance cue that could enable them to identify prospective mates in a noisy chorus environment where the discrimination of individual calls is often difficult. Generally, such redundant signals may facilitate mate choice by enhancing the quality and accuracy of information females receive regarding prospective mates.     

The interrelationships between resource-holding potential,resource-value and reproductive success in territorial males: How much variation can we explain?

A long-standing hypothesis in behavioural ecology posits that males with greater resource-holding potential (RHP) control resource sites deemed more valuable by sexually-receptive females and, thereby, males controlling such sites accrue greater reproductive success (RS). This hypothesis has historically been investigated using three separate but non-mutually exclusive relationships (male RHP vs. resource value, resource value vs. male RS and male RHP vs. RS). The relationships between these three variables are predicted to be strongly positive, however, due to measurement error and biological noise, perfect correlations (r = 1.0) are rare in biology even for well-established relationships. Moreover, the inaccurate identification of either the male trait(s) important to RHP or the resource characteristic sought by females will weaken the observed strength of the relationships. Here, I use meta-analysis to quantitatively describe the general pattern of these relationships in animals. I predict that the relationships between male RHP, resource-value and RS should be significantly positive (male RHP and resource-value should explain a large amount of the variation in male RS). My meta-analysis supports this hypothesis; however, in the best case scenario only ca. 20% of the variation in the response variable was explained. I conclude by identifying areas in which we need to improve our investigations of resource-defence animals and recommending approaches to meet these needs. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The evolution of empty nuptial gifts in a dance fly, Empis snoddyi (Diptera: Empididae): bigger isn't always better

Adaptive female choice is thought to have led to the evolution of nutritionally valuable nuptial gifts in many insect species. However, in several dance fly species, males offer and females accept “empty gifts” with no nutritional value. In the species studied here, Empis snoddyi Steyskal, males produce empty balloons comprised of hundreds of silk bubbles and form mating swarms that females approach to investigate males. Males within the swarm engage in agonistic interactions. The empty balloon has been hypothesized to be an indicator of male condition such that males with larger balloons are predicted to have higher mating success and be more successful in male-male interactions than males with smaller balloons. We examined the role of male body size and balloon size in the context of intersexual and intrasexual selection. We found that neither male body size nor balloon size affected the outcome of pairwise male-male interactions. Using multiple-regression techniques, we found significant linear selection for increasing male body size and decreasing balloon size associated with mating success, a surprising result given a positive relationship between male body size and balloon size. A visualization of selection showed the highest peak of male mating success for larger males with intermediate-size balloons. These results can be explained by a trade-off between long-range attraction of females using large balloons and close-range attraction of females via improved flying efficiency associated with smaller balloons. Both male body size and balloon size are important components in determining male mating success; however, the empty balloon does not appear to play a typical role as a sexually selected ornament. Received: 29 December 1997 / Accepted after revision: 7 October 1998     

Body size preferences in the pot-bellied seahorse <Emphasis Type="Italic">Hippocampus abdominalis:</Emphasis> choosy males and indiscriminate females

Male seahorses (genus Hippocampus) provide all post-fertilization parental care, yet despite high levels of paternal investment, these species have long been thought to have conventional sex roles, with female mate choice and male–male competition. Recent studies of the pot-bellied seahorse (Hippocampus abdominalis) have shown that sex-role reversal occurs in high-density female-biased populations, indicating that male mating preferences may lead to sexual selection on females in this species. Egg size, egg number, and offspring size all correlate positively with female body size in Hippocampus, and by choosing large mating partners, male seahorses may increase their reproductive success. While male brood size is also positively correlated with body size, small H. abdominalis males can carry exceptionally large broods, suggesting that the fecundity benefits of female preference for large partners may be limited. We investigated the importance of body size in reproductive decisions of H. abdominalis, presenting focal individuals of both sexes with potential mating partners of different sizes. Mating preferences were quantified in terms of time spent courting each potential partner. Male seahorses were highly active throughout the mate-choice trials and showed a clear behavioral preference for large partners, while females showed significantly lower levels of activity and equivocal mating preferences. The strong male preferences for large females demonstrated here suggest that sexual selection may act strongly on female body size in wild populations of H. abdominalis, consistent with predictions on the importance of female body size for reproductive output in this species. An erratum to this article can be found at     

Females learn from mothers and males learn from others. The effect of mother and siblings on the development of female mate preferences and male aggression biases in Lake Victoria cichlids, genus Mbipia

While species-assortative behaviour is often observed in sympatrically occurring species, there are few examples where we understand the extent to which development of assortative behaviour is genetically or environmentally determined, for instance, through learning. However, the majority of mate choice theory assumes genetic recognition mechanisms. Knowledge about the development of species recognition is important for our understanding of how closely related species can coexist and how this coexistence may have arisen. The ontogeny of female mate choice, for instance, may critically influence the degree of assortative mating under many circumstances. Also, male assortative aggression behaviour may affect fitness and the possibility for coexistence of two closely related species. Here, we test whether male aggression biases and female mate preferences of two Lake Victoria rock cichlid species, Mbipia mbipi and Mbipia lutea, are affected by experience. With an interspecific cross-fostering experiment, we test for the effect of experience with the phenotype of the mother and that of the siblings on species-assortative mate preferences and aggression biases. We demonstrate that female mate preferences are strongly influenced by learning about their mothers’ phenotype but not by experience with their siblings, despite ample opportunity for interactions. Male aggression biases, in contrast, are affected by experience with siblings but not by learning about their mothers’ phenotype. We suggest that the development of assortative behaviour of females, but not of males, creates favourable conditions for sympatric speciation in Lake Victoria cichlids.     

Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki)

In many sexually dimorphic species adult sexes tend to segregate socially, spatially, or in habitat use. Several hypotheses have been formulated regarding underlying mechanisms. We investigated terrestrial habitat use and sexual segregation in a tropical otariid, the Galápagos sea lion (Zalophus californianus wollebaeki), where most of the hypotheses can be ruled out a priori. Factors relating to thermoregulation and costs of locomotion were of prime importance for habitat use. Habitats directly adjacent to the sea, with simple structured flat surfaces, shade, and tide pools were most frequented, but sexes and age classes differed in their usage patterns. Sexual segregation, both spatial and by habitat was pronounced in the reproductive period (RP), but remained high during the nonreproductive period (NRP). A GLM model of habitat use showed that in both seasons adult males frequented habitat types that adult females and other age classes used much less. Males were most abundant in suboptimal inland habitats, which offered only shade for cooling. Females with newborns differed in habitat use from females with older offspring and lone females. Spatial and habitat segregation are explained most parsimoniously as by-products of social processes, primarily intrasexual competition and female avoidance of male harassment, linked to the polygynous mating system.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Tardy females,impatient males: protandry and divergent selection on arrival date in the two sexes of the barn swallow

Protandry reflects the earlier arrival of males than females to the site of reproduction. Such protandry is hypothesised to arise from sex differences in costs and benefits of early arrival. I investigated temporal patterns of arrival date of male and female barn swallows Hirundo rustica and temporal patterns of selection to test the hypothesis that sex differences in selection account for sex differences in arrival date. Mean arrival date of male barn swallows but not of females advanced during the last 33 years, giving rise to an increasing sex difference in arrival date. Early arrival was favoured by increasingly better survival in males, while females showed an opposite pattern that did not reach significance, although the effect differed between sexes. Early arrival increased fecundity in both sexes equally.The sex difference in viability selection in relation to arrival date changed from positive to negative as the degree of protandry increased in recent years, although there was no similar significant relationship for fecundity selection. Furthermore, sex differences in viability selection in a given year affected the degree of protandry in the following year through differential survival of certain phenotypes over others. Finally, temporal changes in sex difference in viability selection and protandry were related to an increase in the interval between first and second clutches, as the duration of the breeding season increased because of climatic amelioration. These findings suggest that arrival date is under divergent selection in the two sexes, providing a mechanism for the evolution of protandry.     

Mass-dependent take-off ability in wintering great tits (Parus major): comparison of top-ranked adult males and subordinate juvenile females

When birds are attacked by predators, initial take-off is crucial for survival. Theoretical studies have predicted that predation risk in terms of impaired flight ability increases with body weight. However, studies in which attacks were simulated, and within-individual daily changes in body weight were used to test mass-dependent take-off outside migration period, have so far failed to show an effect of mass on velocity. In this field study I compared the mass/velocity relationships of alarmed adult male and juvenile female great tits, Parus major. Fattening strategies differ among members of the dominance-structured basic flocks of wintering great tits, and dominant individuals often carry significantly less amount of fat reserves than subordinates. Since the range of body weight gain/loss is the least among dominant males, it was expected that impaired flight ability is more likely in lower-ranked female great tits. The results show that the birds differed significantly in their daily increase of relative body weight. Average daily weight increase of adult males was 6.2%, while it was 12.2% in juvenile females. Males were faster than females at take-off both at dawn and at dusk. Flight velocity of males did not differ significantly between dawn and dusk, whereas females took off at a significantly lower speed at dusk than at dawn. The results suggest that the larger fat reserves of subordinate females needed to increase their chances of overwinter survival probably place them at increased risk of predation. Electronic Publication     

Fluctuating sexual dimorphism and differential hibernation by sex in a primate, the gray mouse lemur (Microcebus murinus)

The aim of this study was to investigate reproductive strategies and their consequences in gray mouse lemurs (Microcebus murinus), small solitary nocturnal primates endemic to Madagascar. Previous reports of sexual dimorphism in favor of males and females, respectively, a high potential for sperm competition and pheromonal suppression of mating activity among captive males, led us to investigate mechanisms of intrasexual competition in a wild population. Based on 3 years of mark-recapture data, we demonstrate that sexual dimorphism in this species fluctuated annually as a result of independent changes in male and female body mass. Male body mass increased significantly prior to the short annual mating season. Because their testes increased by 100% in the same period and because their canines are not larger than those of females, we suggest that large male size may be advantageous in searching for estrous females and in enabling them to sustain periods of short-term torpor. In contrast to reports from captive colonies, we found no evidence for two morphologically distinct classes of males. Finally, we also show that most adult males are active throughout the cool dry season that precedes the mating season, whereas most adult females hibernate for several months. This is in contrast to other solitary hibernating mammals, where males typically emerge 1–2 weeks before females. Thus, this first extended field study of M.␣murinus clarified previous conflicting reports on sexual dimorphism and male reproductive strategies in this primitive primate by showing that their apparent deviation from predictions of sexual selection theory is brought about by specific environmental conditions which result in sex-specific life history tactics not previously described for mammals. A general conclusion is that sexual selection can operate more strongly on males without resulting in sexual dimorphism because of independent selection on the same traits in females. Received: 6 July 1997 / Accepted after revision: 28 March 1998     

Daring females,devoted males,and reversed sexual size dimorphism in the sand-dwelling spider <Emphasis Type="Italic">Allocosa brasiliensis</Emphasis> (Araneae,Lycosidae)

Sexual selection theory predicts that a higher investment in offspring will turn females into the selective sex, while males will compete for accessing and courting them. However, there are exceptions to the rule. When males present a high reproductive investment, sex roles can reverse from typical patterns, turning males into the choosy sex, while females locate males and initiate courtship. In many spiders, males are smaller than females, wandering in search of sedentary females and maximizing the number of copulations. In the present study, we present findings on the sand-dwelling wolf spider, Allocosa brasiliensis, evidencing a reversal in typical courtship roles reported for the first time in spiders. Males were bigger than females. Females located males and initiated courtship. Copulation always occurred in male burrows and took place mainly in long burrows. Males donated their burrows to the females after copulation, closing the entrance before leaving with female cooperation from inside. Males would provide females with a secure place for ovipositing, being exposed to predation and diminishing their future mating possibilities until constructing a new burrow. The cost of vacating the burrow and losing the refuge in an unpredictable habitat, such as sand dunes, would explain the courtship roles reversal in this spider species. Results turn A. brasiliensis as a promising model for discussing the determinants of sex roles and the pressures that drive their evolution and maintenance. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Heritability of the algal-foraging ability: an indirect benefit of female mate preference for males' carotenoid-based coloration in the guppy, Poecilia reticulata

Several sexual selection theories assume certain benefits of female mate preference. The direct benefit, i.e., the direct contribution from males to their offspring and females, has been well tested empirically. However, the indirect benefit, i.e., the male's genetic contribution to their offspring, has been poorly demonstrated. Female preference for males' carotenoid-based coloration is known in some animals. Since animals must acquire carotenoids through foods, it is often hypothesized that the brightness of the carotenoid-based coloration is a reliable indicator of the male's foraging ability. Hence, females' indirect benefits, such as greater foraging ability in their offspring, through mate preference for the carotenoid-based coloration are assumed. However, the heritability of the foraging ability for foods that serve as carotenoid resources has not been tested. In this study, a maze experiment was performed in guppies (Poecilia reticulata) to examine the heritability of the foraging ability for algae, carotenoid resources in nature. The latency for completing algal-foraging tasks in this experiment showed high individual variation. Heritable estimates of the foraging ability were substantial (h ² = 0.57 – 0.66) and significant, suggesting a genetic contribution to the foraging ability from fathers to their offspring. This result may support the hypothesis that indirect benefits influence the evolution of female choice.     

Sperm competition in the damselfly Enallagma hageni Walsh (Odonata: Coenagrionidae): benefits of multiple mating to males and females

Summary Sperm competition was investigated in the non-territorial damselfly, Enallagma hageni. Using irradiated (sterile) male techniques, the last male to mate was found to fertilize up to 95% of the eggs of the first clutch laid after mating. Dissection of females collected before, during, and after copula showed that a male removes a maximum of 87% of the sperm by volume of a previous mate. These data verify an earlier estimate of lifetime reproductive success in this species which was based on mating success, and suggest that indirect dissection methods offer minimum estimates of sperm precedence. Male E. hageni have ample opportunity to benefit from sperm precedence, since at least 10% of the receptive females encountered had already mated once that day, but still contained complete or partial clutches of eggs. Female E. hageni benefit directly from high sperm precedence because it allows them to exchange matings for guarding service by males during oviposition bouts under water.     

Differential attraction of females to dominant males: Olfactory discrimination and mating preference in the brown lemming (Lemmus trimucronatus)

Summary This study investigated differential attraction of estrous brown lemmings (Lemmus trimucronatus) to conspecific males recently exposed to each other for a 10-min agonistic encounter. In tests conducted 5 min, 1 h and 24 h after agonistic encounters, females preferred the oder of dominant males to that of defeated males when both odors were presented simultaneously in a Y-maze olfactometer. Defeat in an agonistic encounter did not reduce the propensity of male lemmings to initiate sexual behavior. In one-male, one-female tests conducted 5 min after agonistic encounters, dominant males achieved higher mount and thrust scores while defeated males obtained higher scores for attempted mounts. The sexual behavior of dominant and defeated males did not differ significantly in similar tests conducted 1 h and 24 h later. In contrast, females readily mated with dominant males and tended to avoid defeated males in two-male tethering tests conducted 5 min after agonistic encounters. In these tests, females still showed a preference for dominant males 1 h and 24 h after male agonistic encounters.     

Trade-off between fitness components in males of the polygynous butterfly Callophrys xami (Lycaenidae): the effect of multiple mating on longevity

The differential costs of mating paid by males and females influence the nature and strength of sexual selection. In butterflies, males invest a relatively large amount of time and resources in each mating, but male survival costs of mating have not been demonstrated. I present the results of experiments designed to measure the effect of different aspects of mating on male longevity in the polygynous butterfly Callophrys xami. In experiment 1, I compared the longevity of pairs of males that produced similar amounts of spermatophore, but that mated at different rates, a different numbers of times, and that produced spermatophores at different rates, and found that the longevity of ”low-mating-rate” males was not different from that of ”high-mating-rate” males. In experiment 2, the longevity of virgin males was not significantly different from that of multiply mated males. In experiment 3, I used resource-limited males resulting from experimental food limitation of last-instar larvae; resource-limited virgin males lived significantly more days than resource-limited multiply mated males. Since ecological costs of mating (e.g., disease transmission, predation risk) were excluded in the experiment, diminished male longevity was a product of physiological costs of sexual interactions. These results suggest that the cost of ejaculate production is an important cause of longevity reduction when there are resource limitations; however, the role of other possible physiological costs of mating in longevity reduction is still unknown. Received: 21 March 2000 / Accepted: 26 August 2000     

Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance

Although contemporary animal contest theory emphasises the importance of physical asymmetries in resolving disputes, such asymmetries do not obviously settle fights in all groups. Territorial male butterflies, for example, compete via elaborate non-contact aerial interactions in which success is determined by relative persistence. Prior research suggests that the resolution of these contests is not clearly related to physical variables such as body size or energy reserves. However, given that the contests involve elaborate aerial manoeuvres, one long-standing suggestion is that asymmetries in flight performance, and thus flight morphology, may be important. We addressed this hypothesis via a manipulative investigation into the biophysical correlates of contest success in the speckled wood butterfly, Pararge aegeria. This species possesses the ability for significant adaptive phenotypic plasticity in relevant flight morphological parameters. We took advantage of this plasticity to rear 90 individuals of markedly varying flight morphologies, which we then pitted against each other in a semi-controlled experimental fashion. Multiple logistic and lognormal analyses provided little evidence for the relevance of morphological parameters, including relative flight musculature, wing loading and wing aspect ratio (wing length relative to area), to the outcome and/or duration of experimental contests. Instead, we found a positive effect of age upon contest success. Given that ability for high acceleration is strongly linked to variation in these morphological parameters, our findings suggest that flight performance is not a strong determinant of resource-holding potential in this notably territorial butterfly.     

Biological markets: supply and demand determine the effect of partner choice in cooperation,mutualism and mating

The formation of collaborating pairs by individuals belonging to two different classes occurs in the contexts of reproduction and intea-specific cooperation as well as of inter-specific mutualism. There is potential for partner choice and for competition for access to preferred partners in all three contexts. These selective forces have long been recognised as important in sexual selection, but their impact is not yet appreciated in cooperative and mutualistic systems. The formation of partnerships between members of different classes has much in common with the conclusion of trade agreements in human markets with two classes of traders, like producers and consumers, or employers and employees. Similar game-theoretical models can be used to predict the behaviour of rational traders in human markets and the evolutionarily stable strategies used in biological markets. We present a formal model in which the influence of the market mechanism on selection is made explicit. We restrict ourselves to biological markets in which: (1) Individuals do not compete over access to partners in an agonistic manner, but rather by outcompeting each other in those aspects that are preferred by the choosing party. (2) The commodity the partner has to offer cannot be obtained by the use of force, but requires the consent of the partner. These two restrictions ensure a dominant role for partner choice in the formation of partnerships. In a biological market model the decision to cooperate is based on the comparison between the offers of several potential partners, rather than on the behaviour of a single potential partner, as is implicitly assumed in currently accepted models of cooperation. In our example the members of one class A offer a commodity of fixed value in exchange for a commodity of variable value supplied by the other class, B. We show that when the B-class outnumbers the A-class sufficiently and the cost for the A-class to sample the offers of the B-class are low, the choosiness of the A-class will lead to selection for the supply of high value commodities by the B-class (Fig. 3a). Under the same market conditions, but with a high sampling cost this may still be the evolutionariy stable outcome, but another pair of strategies proves to be stable too: relaxed choosiness of class A coupled with low value commodities supplied by class B (Fig. 3b). We give a number of examples of mating, cooperative and mutualistic markets that resemble the low sampling cost situation depicted in Fig. 3a.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999