The role of melanin- and carotenoid-based plumage coloration in nest defence in the Great Tit

Although plumage coloration is recognized to convey valuable information about the bearer's parental abilities, few studies have explored the relationship between coloration and nest defence. In this study in Great Tit Parus major, we analysed the relationship between nest defence and melanin‐ as well as carotenoid‐based plumage coloration, after controlling for ecological variables known to influence nest defence. A principal components analysis was applied to classify birds according to how vigorously they defended the nest, and the intensity of nest defence was tested against plumage coloration. Males with a large black tie defended their nests more vigorously, but no such effect was found for yellow breast coloration. This suggests that melanin‐based coloration in the Great Tit is associated with aggression, including both dominance‐aggression and nest defence, whereas carotenoid‐based coloration is not. The challenge in future studies will be to demonstrate whether females use this trait as an ornament to assess male quality and whether they trade off between the different ornaments a male may exhibit.     

No association between the melanocortin-1 receptor (MC1R) and black belly stripe size variation in the Great Tit Parus major

Capsule The Great Tit Parus major displays a black melanin breast patch stripe (black tie or black belly stripe) which shows great variation and its size correlates with male breeding success, survival and dominance. We investigated for associations between the melanocortin-1 receptor (MC1R) polymorphism, which has an important function in melanin colouration, and the size of the black belly stripe but were unable to detect any polymorphism in this gene. Variation in the size of the melanin-based black belly stripe may therefore be regulated through genetic variation at other genes or via modification of the gene expression inside the melanocortin system and melanogenesis.     

The function of the variation in the breast stripe of the great tit (Parus major)

Many winter flocking bird species possess a plumage colour polymorphism. Two hypotheses have been proposed to explain the function of this variability in plumage structure: the signalling of (1) individual identity (IRH), or (2) social status (SSH). These two hypotheses were tested on winter flocking great tits, a species showing intraspecific variation in its breast stripe plumage. The SSH was considered the most probable, because a close correlation between social status and breast stripe width was found. The SSH was further tested by using dummy great tits bearing breast stripes of different widths. Great tits that possessed a breast stripe wider than that of the dummy approached it aggressively, and those with a narrower behaved submissively. We therefore conclude that the function of the variation in width of the breast stripe plumage is to proclaim the social status of the individual.     

STRUCTURE AND FORAGING PATTERNS OF FLOCKS OF TITS AND ASSOCIATED SPECIES IN AN ENGLISH WOODLAND DURING THE WINTER

Blue Tits were the commonest and most frequent members of mixed-species insectivorous flocks during the winter at Wytham Wood, Oxford. Six common flocking species (Blue Tit, Longtailed Tit, Great Tit, Coal Tit, Marsh Tit and Golderest) made up nearly two-thirds of the birds in the study area. A tentative interspecific social dominance hierarehy was constructed: Great Tit > Blue Tit > Marsh Tit > Coal Tit > Long-tailed Tit and Goldcrest. Blue Tits permitted conspecifics to approach them more closely than did other flock members. Blue Tits most frequently led the flocks, but no more than would be predicted by their abundance. Great Tits, and Marsh Tits and Coal Tits to a lesser extent, defended territories during the winter. Two of the six common species frequented the inner parts of branches (Great Tit, Coal Tit), two the outer parts of branches (Blue Tit, Marsh Tit), and two the twigs (Long-tailed Tit, Goldcrest). Members of each of these species-pairs showed marked differences in height of foraging and/or species of tree frequented. Because of their abundance, the impact of Blue Tits outside of their most highly frequented foraging zones may exceed that of species concentrating in these other zones (e. g., twigs high in trees). The Coal Tit foraged most diversely, the Great Tit least diversely. Species that foraged diversely in one of the three foraging categories (species of tree, substrate, height) usually foraged less diversely than most other species in the other dimensions. Species usually overlapped least in the zones exploited (inner parts of branches, etc.). Great Tits overlapped with other species less than did any other common flock member, and Coal Tits were the next lowest in this regard.     

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.     

The role of pigment based plumage traits in resolving conflicts

The role of melanin ‘badges of status’, in male–male competition has been well‐studied, in contrast, carotenoid based plumage has largely been examined in the context of female mate choice. Recent work has shown that carotenoid signals can also function in male–male competition, although the functions of the two types of signals is currently unclear. Here, we examine the relationships between colouration, dominance and aggression in the crimson finch Neochmia phaeton, a species where males have both conspicuous red carotenoid plumage and a black melanin patch. We examined the importance of carotenoid and melanin based signals in three contexts: 1) among free‐living birds interacting at a feeding station: we found that neither colour signal influenced the outcome of interactions; 2) in staged dyadic contest in captivity: we found that coloration from carotenoid pigments was positively related to the probability of winning a contest, while the size of the melanin plumage patch was not related to winning; and 3) in staged dyadic contests where male plumage colour had been masked: we found that the number of interactions required to determine dominance increased. While the underlying natural plumage colour was still important in these contests, birds with more intense carotenoid colouration were now more likely to lose. These results confirm carotenoid‐based signalling in male–male contests. However this signal is used in conjunction with other factors such as self‐assessment and body condition. Contrary to traditional expectations, the black melanin patch was not found to be important in this context.     

Feather ornaments are dynamic traits in the Great Tit Parus major

In behavioural studies it has been common to quantify plumage colours or ornaments over a range of dates and link them to fitness characteristics without accounting for seasonal changes in these traits. Such changes are likely to be widespread among birds, yet we lack assessments of this variability within individuals. We studied both within‐ and between‐individual temporal changes in Great Tit Parus major ornaments, specifically the melanin‐based black breast stripe and the pigment‐free white cheek patch. During the non‐breeding season both ornaments varied. In juveniles and adult females, the area of the breast stripe first rose and then, from near the end of December, decreased. In adult males there was a linear decrease. In the cheek patch, the irregularity of the cheek borders showed either a linear (adults) or a non‐linear (juveniles) increase as the season progressed. In individuals repeatedly sampled during the same winter, the decrease in the size of the breast stripe was larger for males than females and there was an overall decrease in the regularity of the cheek borders. There was no relationship between the size of the breast stripe and the white cheek patch irregularities or the cheek patch area. These results imply that more attention should be paid to quantification, within individuals, of the components of expression of phenotypic traits. In addition, we suggest that further research should focus on explaining the causes and functions of ornament change.     

Plumage coloration can be perceived as a multiple condition‐dependent signal by Great Tits Parus major

The evolution of multiple signals can be explained because they enhance the perception of a general message by recipients. Plumage coloration frequently acts as a condition‐dependent signal, so that species displaying different colour patches have the potential to transmit information on condition through a multiple signalling system. The Great Tit Parus major exhibits plumage colour patches generated by the main types of colour production, some of which, particularly those based on melanins and carotenoids, are known to be related to body condition. However, the colour expressions of all of the different colour types have never been investigated simultaneously to determine whether they function as multiple signals of condition. In addition, visual perception models have never been applied to a multiple signalling system in a wild population of birds. Here I present information that links body condition with the colour expression of almost all of the different plumage patches of male Great Tits captured during the winter. Birds in better condition had greater reflectance values at short wavelengths in all plumage patches, and this was especially so in the white (i.e. structural) colour of the cheeks. Plumage colour characteristics were calculated by means of avian visual models, suggesting that Great Tits have the capacity to perceive information contained in the plumage coloration of conspecifics. These results show that short‐wavelength reflectance has great potential to transmit biologically significant information on the body condition of birds, even in achromatic plumage patches.     

Cryptic sexual dichromatism occurs across multiple types of plumage in the Green-backed Tit Parus monticolus

Several recent studies have found instances of cryptic sexual dichromatism within avian taxa. Although this dichromatism has been found in plumage produced through a variety of proximate mechanisms, little is known about how dichromatism varies across these types of plumage within a single species. We used a reflectance spectrometer to measure colour within the Green-backed Tit Parus monticolus , a species which displays multiple types of pigment and structural colours. We found significant differences in spectral measurements corresponding to hue, chroma, and brightness between male and female carotenoid, melanin, structural white, grey and structural blue plumage. The only plumage that did not appear to show sexual dichromatism was the olive plumage of the back. These findings suggest that the mechanism(s) producing cryptic dichromatism in the Green-backed Tit are non-specific and act across multiple types of plumage, rather than within a single type, such as carotenoid-based or structurally produced.     

Carotenoid plumage hue and chroma signal different aspects of individual and habitat quality in tits

We hypothesized that Blue Tits Cyanistes caeruleus and Great Tits Parus major from low quality habitat (small woods) would have less yellow ventral plumage than those from high quality habitat (large woods) because they moult faster and/or their diet contains fewer carotenoids. They moult faster because they moult later in the season and are subject to more rapidly shortening daylengths. We tested this using a database of the plumage coloration (chroma, hue and lightness) of birds breeding in woods of different sizes, by manipulating the speed of moult in captive Blue Tits, and by counting the abundance and size of caterpillars (the major source of dietary carotenoids) in the diet of nestlings. In accordance with our hypothesis, juveniles of both species (which moult about three weeks later than adults) were about 8% less saturated in colour (lower chroma) than adults, but there was no significant difference in chroma between habitats. However, both species did differ significantly in hue between large and small woods. Blue Tits forced to moult faster in captivity, at a rate similar to that caused by a month's delay in the start of moult, had yellow flank feathers that were 32% less saturated in colour than those allowed to moult more slowly. Blue Tit nestlings in large woods consumed 47% more caterpillar flesh (per gram of faecal material voided) than those in small woods, and Great Tit pulli 81% more. When habitat effects were controlled for in ANOVAs, Blue Tits mated assortatively on the basis of flank hue and Great Tits on the basis of flank lightness. Flank colour therefore has the capacity to provide information about the potential quality of both habitats, and individual birds, to potential colonists and sexual partners.     

Pollination of the Crown ImperialFritillaria imperialis by Great TitsParus major

Visitations of flowers of the Crown Imperial by Great Tits at two different locations in Hessen, Germany, in 1990 and 1993 are described. The observations prove that flowers were visited because of the nectar; most likely pollination occurs during this visits. Similar reports from Europe are discussed. The significance of specific interactions including ecological generalists such as the Great Tit is considered in the context of the phylogenetic reconstruction of the development of co-adaptive syndroms.

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Zusammenfassung Beobachtungen von Kohlmeisen (Parus major) an Blütenständen der Kaiserkrone (Fritillaria imperialis) in Hessen zeigen, daß der Blütenbesuch des Nektars wegen erfolgte und daß dadurch mit größter Wahrscheinlichkeit eine Bestäubung der Pflanzen erreicht wird. Ähnliche Fälle aus Europa werden diskutiert. Die Bedeutung spezifischer Interaktionen unter Beteiligung ökologischer Generalisten wie der Kohlmeise für die Rekonstruktion der phylogenetischen Entstehung co-adaptiver Syndrome wird erörtert.

     

Seasonal changes in carotenoid- and melanin-based plumage coloration in the Great Tit Parus major

Plumage coloration plays an important role as a signal of individual quality. We studied plumage coloration in two Great Tit Parus major populations to test the hypothesis that coloration changes during the year without moult. As expected, the coloration of 'yellow' and 'dark green' areas in the breast and back changed with time but not the colour characteristics of the 'black' crown. This suggests that colours based on carotenoids could be more exposed to abrasion or fading than melanin-derived colours, although other factors could affect the patterns of change found. Our results show that effects of colour alteration could be important to the understanding of the potential for reliable signalling of different colours. Seasonal changes in plumage colour should be considered in future analysis of intraspecific variation in coloration.     

Nestbox orientation: a species-specific influence on occupation and breeding success in woodland passerines

Capsule Nestbox orientation has species-specific influences on nestbox occupation and breeding success for woodland passerines.

Aims To determine if nestbox orientation had an influence upon nestbox selection or breeding success for three co-occurring woodland passerines.

Methods We analysed 15 consecutive years of breeding data (1990–2004) from 295 nestboxes in the UK using circular statistical analyses to examine the influence of orientation upon nestbox occupation and breeding success for three species, Blue Tit Cyanistes caeruleus, Great Tit Parus major and Pied Flycatcher Ficedula hypoleuca.

Results The three species used nestboxes of all orientations during the 15-year period. The frequency of nestbox occupation by Great Tits correlated with orientation (the mean number of nests in boxes oriented south-southwest was lower than the mean number of nests in boxes facing other directions). There was no such relationship for Blue Tits or Pied Flycatchers. Nestbox orientation influenced the breeding success of Pied Flycatcher (the mean number of young to fledge from boxes oriented south-southwest was lower than from boxes facing other directions). There was no such relationship for Blue or Great Tits.

Conclusion Nestbox orientation can be an important influence on occupation and breeding success, but this differed between species. Intriguingly, although the directionality reduced nestbox occupation (Great Tit) and breeding success (Pied Flycatcher) was the same (south-southwest), there was a disparity in the influence of orientation for Great Tit (orientation influenced the frequency of occupation but not success) and Pied Flycatcher (orientation did not influence occupation but did affect success). We discuss these disparities, considering the possible influences of mating strategy, breeding phenology, nestbox microclimate and offspring quality.     

破碎化次生林斑块面积及斑块隔离对大山雀繁殖成功的影响

研究了破碎化山地次生林中斑块面积及斑块隔离度对大山雀（Parus major)繁殖成功的影响,运用GPS定位系统确定了18块大,中,小3种类型的斑块及对照样点,观测了大山雀产第一枚卵时间,窝卵数,平均卵重,出雏量及雏鸟出飞量等生态指标,结果表明,斑块隔离度对大山雀繁殖成功没有影响,1999－2000年两年中,大山雀在连续分布次生林中的产卵时间平均早于各斑块中的产卵时间7．2d,各斑块间的产卵时间差异较小,连续分布次生林和较大面积斑块内的大山雀窝卵数略高于中,小面积斑块内的窝卵数,连续分布生林中的平均卵重最大,斑块面积对出雏量及雏鸟不量没有影响,中,小面积斑块内的巢损失率较高,最主要的原因是巢址竞争较激烈。     

Evolutionarily stable transition rates in a stage-structured model. An application to the analysis of size distributions of badges of social status

This paper deals with the adaptive dynamics associated to a hierarchical non-linear discrete population model with a general transition matrix. In the model, individuals are categorized into n dominance classes, newborns lie in the subordinate class, and it is considered as evolutionary trait a vector eta of probabilities of transition among classes. For this trait, we obtain the evolutionary singular strategy and prove its neutral evolutionary stability. Finally, we obtain conditions for the invading potential of such a strategy, which is sufficient for the convergence stability of the latter. With the help of the previous results, we provide an explanation for the bimodal distribution of badges of status observed in the Siskin (Carduelis spinus). In the Siskin, as in several bird species, patches of pigmented plumage signal the dominance status of the bearer to opponents, and central to the discussion on the evolution of status signalling is the understanding of which should be the frequency distribution of badge sizes. Though some simple verbal models predicted a bimodal distribution, up to now most species display normal distributions and bimodality has only been described for the Siskin. In this paper, we give conditions leading to one of these two distributions in terms of the survival, fecundity and aggression rates in each dominance class.     

Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones

Pseudopodoces humilis (Hume's Ground-Jay) is a small passerine bird that inhabits the high rocky steppes of the Tibetan (Qinghai–Xizang) Plateau. Although it was long classified as a small species of ground jay ( Podoces ), two previous anatomical studies cast doubt on its assignment to the Corvidae (crows and jays). We studied the evolutionary relationships of Pseudopodoces using three independent datasets drawn from comparative osteology, the nuclear c- myc gene, and the mitochondrial cytochrome b gene. All three datasets agree on the placement of Pseudopodoces in the family Paridae (tits and chickadees). The cytochrome b data further suggest that Pseudopodoces may be closest to the Great Tit Parus major species group. Pseudopodoces is the only species of parid whose distribution is limited to treeless terrain. Its evolutionary relationships were long obscured by adaptations to open habitat, including pale, cryptic plumage; a long, decurved bill for probing in crevices among rocks or in the ground; and long legs for terrestrial locomotion. Despite these accommodations to a novel adaptive zone, its evolutionary affinity with the Paridae is clearly expressed in comparative osteology and genetics, and is supported by its habit of nesting in cavities.     

The evolution of aggression: can selection generate variability?

Three models--the war of attrition, the size game and the badges of dominance game--are described, in which natural selection can maintain genetic variability for aggression. The models differ in whether or not the traits that settle contests are costly in contexts other than fighting, and also in whether signals are used. It is concluded that contests will be settled by non-costly traits only if the value of the contested resource is small relative to the cost of fighting, and that 'honest' signalling of aggressiveness is stable only if individuals giving signals that are inconsistent with their behaviour suffer costs. The literature on 'badges of dominance' in birds is reviewed. New data on great tits, greenfinches and corn buntings show that there is plumage variability within age and sex that sometimes serves to settle contests, and that, in the first two species but not the third, the badges are uncorrelated with size, and settle contests only over trivial resources.     

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.     

Genetically‐based behavioural morph affects stopover refuelling performance in white‐throated sparrows Zonotrichia albicollis

Intraspecific competition can influence refuelling at migration stopover sites. White‐throated sparrows Zonotrichia albicollis have genetically‐determined plumage morphs that differ in dominance behaviour and competitive abilities. This study examines the effects of plumage morph, sex and age, three likely indicators of competitive ability, on fall migration timing, body composition, and refuelling rates during stopover at Long Point, Ontario. We used quantitative magnetic resonance analysis and plasma metabolite profiling to determine body composition and refuelling rates, respectively. We determined sex and plumage morph genetically. Competitive ability did not influence migration timing. Controlling for structural size, males had larger lean mass than females, but we found no differences in body fat or lean mass between plumage morphs. Plasma metabolite concentrations indicated that the aggressive white‐stripe morph refuelled faster than the less aggressive tan‐stripe morph, though there were no differences among sex and age groups. We suggest that increased refuelling rates did not result in increased fat or lean mass because 1) individuals categorised as less competitive had longer stopover durations to compensate for slower refuelling rates; or 2) costs of behaving more competitively offset gains from faster refuelling rates.