Manipulating badges of status only fools strangers

Conflict is risky, but mechanisms that allow animals to assess dominance status without aggression can reduce such costs. Two different mechanisms of competitor assessment are expected to evolve in different contexts: badges of status are expected in larger, anonymous groups, whereas individual recognition is feasible in small, stable groups. However, both mechanisms may be important when social interactions occur both within and across stable social groups. We manipulated plumage in golden‐crowned sparrows (Zonotrichia atricapilla) and found that two known badges of status – gold and black head plumage patch sizes – independently affect dominance among strangers but manipulations had no effect on dominance among familiar flockmates. Moreover, familiar flockmates showed less aggression and increased foraging relative to strangers. Our study provides clear experimental evidence that social recognition affects badge function, and suggests that variation in social contexts maintains coexistence and context‐dependent use of these two dominance resolution mechanisms.     

BADGE SIZE,PHENOTYPIC QUALITY,AND REPRODUCTIVE SUCCESS IN THE HOUSE SPARROW: A STUDY ON HONEST ADVERTISEMENT

Honest signaling theory suggests that advertising traits must be costly to their bearer; thus, only individuals of high phenotypic quality can exhibit maximal expression of these traits. Males of the sexually dichromatic house sparrow, Passer domesticus, have a black throat patch that functions as a badge of status. I investigated whether badge size honestly shows phenotypic quality. Badge size increases with age and decreases with advancing fledging date in yearling males; thus, badge size was larger in older individuals even though age differences were small. Badge size also increased with physical condition independent of age. These results indicate that badge size functions as an honest signal, possibly because there are costs involved in its production. I also found that males with enlarged badges acquired more nest sites than either control males or males with reduced badges. However, males with enlarged badges possessing a nest site raised fewer fledglings per year than did males with reduced badges, suggesting that cheating has no selective benefit. Further studies that accurately measure the energy expenditure allocated to badge production and that quantify additional fitness components are needed to clarify how reliable badges are maintained.     

Social Control and Physiological Cost of Cheating in Status Signalling Male House Sparrows (Passer domesticus)

Flock-forming passerines often use plumage characteristics to signal their social dominance. While the benefits to signal dominance seem obvious, costs associated with status signalling are ambiguous. The social control hypothesis predicts that individuals of high social status – with large badges – are involved in more social interactions with individuals of similar badge size. Cheaters are therefore exposed to increased risk of fighting with high quality individuals and the costs associated with enhanced fights with dominant males are supposed to outweigh the benefits of cheating. We tested the social control hypothesis in male house sparrows ( Passer domesticus ), by observing social interactions in captive flocks and determining dominance relationships. Two low status individuals within each flock had the size of their badge experimentally increased and the interactions involving experimental and control birds were recorded. We also assessed the potential physiological cost of cheating in terms of enhanced levels of the stress hormone, corticosterone. Dominance was significantly positively correlated with badge size, but not with other morphological traits. We found little support for the social control hypothesis. Birds did not have significantly more interactions with individuals of similar badge size, before the manipulation. Similarly, after the experimental increase in badge size, experimental birds did not tend to have more encounters with large-badged males. Experimental birds with enlarged badges won more fights compared with prior to the manipulation, suggesting that badge size is used as a signal of social dominance even in small and stable flocks. Finally, corticosterone levels in the blood did not increase significantly after the manipulation of badge size, suggesting that there is no measurable cost, resulting from stress, in cheaters.     

An investigation of mate choice based on manipulation of multiple ornaments in Kentish plovers

Many animals have multiple sexual ornaments, a fact variously explained as signalling of multiple attributes, or nonadaptive retention of now redundant, but previously informative, signals. Despite the widespread occurrence of multiple ornaments, and the theoretical interest in how they are maintained by selection, there have been few experimental studies of the phenomenon. We investigated the role of two ornaments, each plausibly signalling different male attributes, in attracting a new mate in the Kentish plover, Charadrius alexandrinus. Previously we have shown that male Kentish plovers vary in how long they take to acquire a new mate, and we hypothesized that this variation may relate to their attractiveness or parental ability. We created single males by removing their mate and clutch, and then manipulated both their badge size (a presumed signal of either their genetic quality or their dominance status and hence defensive abilities) and the length of their flank feathers (a presumed signal of their parental quality in incubation) in a 2×2 factorial design. We found no difference in remating times between manipulated and control males. Furthermore, neither body size nor body condition of males was related to their remating times, although males with enlarged badges spent less time fighting than control males. Taken together, our results suggest that female Kentish plovers do not use either badge size or the length of flank feathers as cues in their mate choice decisions. However, badge size may influence male-male competition.     

Experimentally increased badge size increases male competition and reduces male parental care in the collared flycatcher

Experimental enlargement of sexually selected traits that are energetically cheap to produce is expected to reveal costs resulting from increased risk of predation or social competition. Given a trade-off between sexually selected traits and life history traits such costs may be expected to affect not only the males themselves but also their offspring. In this study I manipulated the size of the forehead patch, a sexually selected trait that functions as a badge of status in male collared flycatchersFicedula albicollis). First, I found that a male''s likelihood to establish a breeding territory with respect to his original badge size was affected by the treatment such that old males (older than or equal to two years) with relatively small original badges enjoyed an increased likelihood of establishing a breeding territory while young males (yearlings) suffered a reduced likelihood of establishment when their badges were enlarged as compared to unchanged. Second, young males with enlarged badges that were able to establish a territory fed their nestlings less in relation to their females compared to the control males. However, the females adjusted their parental effort to such an extent that no significant differences were observed in total feeding rate nor in reproductive success between the two groups of males. These results suggest that experimentally enlarged badge size in the collared flycatcher may result in increased male competition and that males have to trade their effort spent in male contest against their parental effort.     

Social runaway: Fisherian elaboration (or reduction) of socially selected traits via indirect genetic effects

Our understanding of the evolutionary stability of socially selected traits is dominated by sexual selection models originating with R. A. Fisher, in which genetic covariance arising through assortative mating can trigger exponential, runaway trait evolution. To examine whether nonreproductive, socially selected traits experience similar dynamics—social runaway—when assortative mating does not automatically generate a covariance, we modeled the evolution of socially selected badge and donation phenotypes incorporating indirect genetic effects (IGEs) arising from the social environment. We establish a social runaway criterion based on the interaction coefficient, ψ , which describes social effects on badge and donation traits. Our models make several predictions. (1) IGEs can drive the original evolution of altruistic interactions that depend on receiver badges. (2) Donation traits are more likely to be susceptible to IGEs than badge traits. (3) Runaway dynamics in nonsexual, social contexts can occur in the absence of a genetic covariance. (4) Traits elaborated by social runaway are more likely to involve reciprocal, but nonsymmetrical, social plasticity. Models incorporating plasticity to the social environment via IGEs illustrate conditions favoring social runaway, describe a mechanism underlying the origins of costly traits, such as altruism, and support a fundamental role for phenotypic plasticity in rapid social evolution.     

Different Reproductive Tactics in House Sparrows Signalled by Badge Size: Is There a Benefit to Being Average?

Sexual selection models usually predict directional selection for ornamental traits because of intra‐ as well as inter‐sexual selection. Animals frequently face reproductive trade‐offs, such as between mating and parental effort. Provided that both are essential and have opposite effects on ornament expression, we may however not necessarily expect directional selection for ornament size. The house sparrow is an ideal species to study such a trade‐off, as the size of the male ornament, the black throat badge, seems to be inversely related to mating and parental effort. It has been suggested that large‐badged males invest more in female attraction and territory defence, while small‐badged males may invest more in parental care. In a nest‐box study, we show that females started to breed earliest and produced the largest clutches when mated to males with average‐sized badges that invested in paternal care more than other males. These results are discussed in view of inter‐ as well intra‐sexual selection. Overall, average‐badged males experienced the highest hatching failures, their chicks were in the poorest physical condition and they did not fledge more chicks than other males. It is therefore unlikely that the mating advantages that we observed could by themselves lead to stabilizing selection for badge size. Our results rather suggest that badge size in male house sparrows signals different reproductive tactics, which are adapted flexibly according to their physical condition and socio‐ecological situations.     

Honest signalling,dominance hierarchies and body condition in House Sparrows Passer domesticus (Aves: Passeriformes) during acute coccidiosis

Parasitic infections may change the equilibrium between the costs and benefits of an animal for maintaining its status in a social group. Consequently, parasites may influence the social status of an animal in a group. The present study investigated whether acute infection with Isospora spp. has any effect on the social relationships (e.g. dominance hierarchy) of male house sparrows and how the infection influences their behaviour, immune status, and body condition. Furthermore, the study allowed us to examine how important the ‘badge of dominance’ is with respect to maintaining social status even when the actual condition is changing as a result of infection. The results obtained showed that an acute infection leads to changes in the dominance hierarchy of a social group and that body mass losses of birds depend on the achieved hierarchy status. A positive correlation between the badge size and male aggressiveness was only found during acute infection. In addition, we also found a relationship between cell‐mediated immune response and male aggressiveness during acute infection. This suggests that male badge size is not sufficient to maintain a given dominance position. On the other hand, badge size, a signal developed during the moult, appears to remain an informative and ‘honest’ signal several months later, reflecting the energy reserves of a bird faced with a demanding stressful situation such as acute infection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 718–726.     

Badge size, paternity assurance behaviours and paternity losses in male house sparrows

Male quality may influence both the outcome of sperm competition and female faithfulness. In male house sparrows Passer domesticus , the size of the black throat patch (badge) signals dominance and perhaps attractiveness. So far, however, no study has reported any significant relationships between badge size, paternity and paternity assurance behaviours in this species. We found that the time mates spent together at the nest was positively correlated with badge size. Furthermore, although paternity losses were influenced by both the time spent at the nest and within-pair copulation frequency, we found no relationship between copulation rate and badge size. It seems therefore that copulation frequency served as a paternity assurance behaviour, whereas the time mates stayed together at the nest may have reflected male attractiveness. Alternatively, females may have decided to stay with large-badged males because they were better able to protect them from harassment by strange males. We also found that paternity losses were related to male badge size; average-badged males cuckolded were more often than males with smaller or larger badges. We suggest that average-badged males suffered higher paternity losses because they had different time allocation strategies than other males.     

Testosterone levels in relation to size and UV reflectance of achromatic plumage traits of female pied flycatchers

In a substantial number of species, females show some development of secondary sexual characters. These traits can function as signals of individual phenotypic or genetic qualities and status to conspecifics. Individuals may benefit potentially from expressing signals or badges of status if they are reliable and honest signals of individual quality. In many species, badge sizes have been shown to correlate with dominance rank, which may be mediated by testosterone (T) levels. Here, we explored geographic variation in the size and properties of the white wing patch of female pied flycatchers Ficedula hypoleuca and its relation to circulating T levels in three populations (two southern populations in central Spain and a northern population in Finland). Furthermore, we aimed at detecting if the size of the white wing patch and its ultraviolet (UV) reflectance indicate individual quality. We found that females in Spain had larger, brighter and more UV reflecting wing patches than those in Finland. Females with higher UV reflectance and larger primary white patches bred earlier. Younger females and females with larger primary white wing patches showed higher T levels. In contrast, higher values of UV reflectance in feathers from these patches were associated with low T levels. Despite genetic differentiation and differences in trait expression between populations, female pied flycatchers from different populations may converge and use the size of white wing patches to signal their T levels and thereby their social dominance.     

The role of testosterone in bib size determination in the male house sparrow Passer domesticus,is age dependent

Secondary sexual signals are thought to indicate individual quality. In order to understand the evolutionary pressures that give rise to such traits it is important to understand the physiological mechanisms underlying their production. The black bib of the house sparrow Passer domesticus is known to function as a badge of social status in males. Past studies have found that the size of the bib in older males is determined, at least partly, by the androgen testosterone. The immunocompetence handicap hypothesis suggests that testosterone has a key role in maintaining honest signalling – it is both involved in the development or expression of sexual signals and is immunosuppressive. In this paper we test experimentally two hypo theses relating to bib size development, whether 1) testosterone is only immunosuppressive in conditions where the natural feedback loop from the testes has been removed, and 2) testosterone is, in addition to influencing the bib size of older males, responsible for the size of the bib in juvenile sparrows. In the first experiment we found that exogenous testosterone administered to intact males during the winter (when LH and FSH levels are very low and were not artificially increased by castration) caused significant immunosuppression, albeit in interaction with the stress hormone corticosterone. Second, we found that exogenous testosterone administration in castrated fledgling male house sparrows had no effect on subsequent post‐juvenile moult bib size relative to controls. Our results suggest that in some circumstances testosterone can be immunosuppressive, but that its role in bib size determination is age‐dependent.     

Bimorphism in Male Verreaux’s Sifaka in the Kirindy Forest of Madagascar

Male primates in species with pronounced secondary sexual adornments can exhibit reversible or irreversible bimorphism, i.e., striking variation in the degree to which males express the adornments. Verreaux’s sifaka (Propithecus verreauxi verreauxi) use scent marking as a form of communication and exhibit sex differences in scent glands. Some males exhibit a pronounced brown staining around their sternal gland, whereas others do not. We studied morphological and behavioral characteristics of males in 6 social groups in Kirindy Forest, Madagascar, from November 2000 to March 2002 to evaluate the hypotheses that the bimorphism in male sifaka chest status represents alternative mating tactics and is a badge of status. Males are clearly divided into 2 categories: clean and stained chests, with rare, but informative, intermediate males. The chest staining probably results from the males scent marking with their sternal glands, because stained-chested males scent marked significantly more often than clean-chested males. Though sample sizes are small, chest status did not appear to depend on body size. Chest status is reversible and related to dominance rank. In each group, only 1 male, the dominant, was stained-chested, whereas all other (subordinate) males were clean-chested. These findings suggest that stained chests are visual and olfactory signals of dominance rank and that clean chests signal lack of competitive intent. Thus, this bimorphism may reflect alternative mating tactics used by males to maximize their reproductive success based upon their social environment.     

Group size structure affects patterns of aggression in larval salamanders

The potential importance of intrapopulation phenotypic variabilityto population-level ecology has been demonstrated in both theoreticaland field studies. One way to connect individuals to the dynamicsof populations they compose is to study behavioral response(an individual-level characteristic) to variability in conspecificphenotypes (a population-level characteristic). We examinedeffects of variation in size of individuals on patterns of aggressionin larval tiger salamanders (Ambystoma tigrinum nebulosum) byobserving aggressive behavior in groups of three larvae in alaboratory experment. We assessed effects of variability insize of conspecifics independently of mean larval size and larvaldensity Overall levels of aggression were generally higher ingroups in which all individuals were similprly sized than ingroups of variably sized individuals. Medium-sized individualsexhibited significantly higher levels of aggression and wereattacked significantly more often when in groups consistingonly of similarly sized larvae as compared to groups composedof larvae representing a wider range of body sizes. Activitylevels of larvae were also generally lower when all individualswere the same size, resulting in a negative correlation betweenactivity and levels of iggressititi. These results suggest thatgioups of similarly sized individuals experience a more aggressivesocial environment than groups of variably sized individuals,and they suggest a promising avenue of research for connectingindividual behavioral and physiological responses to size structure(phenotypic variability) with population dynamics.     

Multiple Cues in Status Signalling: The Role of Wingbars in Aggressive Interactions of Male House Sparrows

During aggressive interactions, animals may signal their competitive ability by various ornaments referred to as badges of status. The use of a single badge predicting dominance rank occurs in many vertebrate species. However, animals often display multiple ornaments that may convey information about either different or the same aspects of the signaller's quality, or alternatively, may serve as signal amplifiers. We observed the fighting behaviour of male house sparrows in two captive flocks to investigate whether they may use multiple cues in status signalling during aggressive interactions. Beside the status‐signalling bib, male sparrows possess a conspicuous white wingbar that they often display upon aggressive encounters. We tested whether bib size and the wingbar's conspicuousness (i.e. its achromatic contrast with the neighbouring dark feathers) or its area predicted success in various aspects of fighting. We found that bib size strongly predicted overall fighting success (i.e. proportion of fights won) and defence success (i.e. proportion of successful defences out of all attacks received). Wingbar conspicuousness was positively related to defence success after controlling for the effect of bib size in multivariate analyses. Furthermore, displaying the wings also tended to improve the birds’ success in defence but not in attack. Wingbar area was unrelated to any measured aspect of fighting ability. We suggest that bib size and wingbar conspicuousness may convey multiple messages on fighting abilities, specifically on overall aggressiveness and defending potential, respectively. Alternatively, wingbars may serve as amplifiers for the wing displays of aggressive motivation. Thus, male sparrows may use multiple cues in assessing the competitive ability of opponents during social interactions.     

Differential selection according to the degree of cheating in a status signal

The maintenance of honesty in a badge-of-status system is not fully understood, despite numerous empirical and theoretical studies. Our experiment examined the relationship between a status signal and winter survival, and the long-term costs of cheating, by manipulating badge size in male house sparrows, Passer domesticus. The effect of badge-size manipulation on survival was complex owing to the significant interactions between the treatments and original (natural) badge size, and between the treatments and age classes (yearlings and older birds). Nevertheless, in the experimental (badge-enlargement) group, males with originally large badges had increased winter survival, while males with originally small badges had decreased survival. This indicates that differential selection can act on a trait according to the degree of cheating.     

Uropygial gland size,feather holes and moult performance in the House Sparrow Passer domesticus

Feather holes represent damage to the plumage of birds and are correlated with delayed moult. Uropygial gland size is negatively correlated with feather holes. Consequently, it was predicted that birds with smaller uropygial glands would have more feather holes, and that this would affect moult performance. I examined this prediction in the House Sparrow Passer domesticus. Individuals with smaller uropygial glands had more feather holes, and those with more feather holes moulted later and faster. Therefore, uropygial gland size seemed to affect moult performance via its effect on feather holes. Uropygial gland size may have a positive effect on plumage quality, through a negative effect on feather holes, and therefore on moult timing and speed.     

Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

Environmental influence and cohort effects in a sexual ornament in the house sparrow, Passer domesticus

Presence of phenotypic variation is necessary for selection to occur, yet processes affecting variation in sexually selected characters in natural populations are poorly understood. Here we examine whether variation in a sexual ornament (badge size) of male house sparrows ( Passer domesticus ) is dependent on individual variation in the conditions during early ontogenetic stages, and whether this variation and the population-wide effects of external variables such as weather or population size jointly will generate consistent differences among cohorts later in life. Variation in badge size was independent of adult body size, whereas heavier fledglings and fledglings in good body condition developed smaller visible badges as adults. Furthermore, strong cohort-effects were present, caused by a combined effect of density-dependence and weather during the early development in the moulting period and autumn after hatching. Thus, badge size is an environmentally-dependent trait in house sparrows, and likely to be under the influence of both natural and sexual selection.     

Comparison of EPR occupational lifetime external dose assessments for Mayak nuclear workers and film badge dose data

The Mayak worker cohort is one of the major sources of information on health risks due to protracted exposures to plutonium and external ionizing radiation. Electron paramagnetic resonance (EPR) measurements in tooth enamel in combination with personal dose monitoring can help to improve external dose assessment for this cohort. Here, the occupational lifetime external exposure was evaluated individually for 44 nuclear workers of three plants of the Mayak Production Association by EPR measurements of absorbed doses in collected tooth enamel samples. Analysis included consideration of individual background doses in enamel and dose conversion coefficients specific for photon spectra at selected work areas. As a control, background doses were assessed for various age groups by EPR measurements on teeth from non-occupationally exposed Ozyorsk residents. Differences in occupational lifetime doses estimated from the film badges and from enamel for the Mayak workers were found to depend on the type of film badge and the selected plant. For those who worked at the radiochemical processing plant and who were monitored with IFK film badges, the dose was on average 570 mGy larger than estimated from the EPR measurements. However, the average difference was found to be only −4 and 6 mGy for those who were monitored with IFKU film badges and worked at the reactor and the isotope production plant respectively. The discrepancies observed in the dose estimates are attributed to a bias in film badge evaluation.N. El-Faramawy: On leave from Department of Physics, Faculty of Science, Ain Shams University, 65511 Abbassia, Cairo, Egypt.     

Group size versus territory size in group-living badgers: a large-sample field test of the Resource Dispersion Hypothesis

Badgers ( Meles meles ) have been the focus for the development of a pervasive model of social grouping behaviour, relevant to a number of carnivore species and other taxonomic groups – the Resource Dispersion Hypothesis (RDH). The RDH hypothesises that the dispersion and richness of resources in the environment provide a passive mechanism for the formation of groups, even without any direct benefits of group living. However, few studies have tested the RDH in the field. The principal prediction is that, as opposed to enlargement of territory sizes to accommodate more members, territory size (TS) is independent of group size (GS). Instead, TS is determined by the spatial dispersion of resources, while GS is independently determined by the richness of those resources. However, these predictions provide only weak correlative tests, especially in non-experimental field studies. The first predicts an absence of correlation and is therefore prone to Type II error, especially given the small sample sizes and errors in estimating TS and GS of mammals in the field. We tested for independence of territory size and group size in all years with available data since the beginning of the long-term badger study in Wytham Woods in 1974. We used two methods of TS estimation, a sequential Bonferroni technique to adjust for multiple inference tests, a combined analysis and an analysis with pooled data. This prediction of the RDH could not be rejected on the basis of any of these analyses. Given this evidence that other processes are independently determining group size and territory size, further predictions of the RDH will be worth investigating in considerable detail.