Feather Growth Rate Research Articles

Molt–breeding overlap alters molt dynamics and behavior in zebra finches,Taeniopygia guttata castanotis

Costly events in the life history cycle of organisms such as reproduction, migration and pelage/plumage replacement are typically separated in time to maximize their outcome. Such temporal separation is thought to be necessitated by energetical trade-offs, and mediated through physiological processes. However, certain species, such as tropical birds, are able to overlap two costly life history stages: reproduction and feather replacement. It has remained unclear how both events progress when they co-occur over extended periods of time. Here we determined the consequences and potential costs of such overlap by comparing molt and behavioral patterns in both sexes of captive zebra finches (Taeniopygia guttata castanotis) that were solely molting or were overlapping breeding and molt. Individuals overlapping the early stages of breeding with molt showed a roughly 40% decrease in the growth rate of individual feathers compared with birds that were molting but not breeding. Further, individuals that overlapped breeding and molt tended to molt fewer feathers simultaneously and exhibited longer intervals between shedding consecutive feathers on the tail or the same wing as well as delays in shedding corresponding flight feathers on opposite sides. Overlapping individuals also altered their time budgets: they devoted more than twice the time to feeding while halving the time spent for feather care in comparison to molt-only individuals. These data provide experimental support for the previously untested hypothesis that when molt and reproduction overlap in time, feather replacement will occur at a slower and less intense rate. There were no sex differences in any of the variables assessed, except for a tendency in females to decline body condition more strongly over time during the overlap than males. Our data indicate the existence of major consequences of overlapping breeding and molt, manifested in changes in both molt dynamics and time budgets of both sexes. It is likely that under harsher conditions in natural environments such consequences will be more severe and may result in fitness consequences.

Many quantitative trait loci for feather growth in an F2 broiler × layer cross collocate with body weight loci

1. A genome-wide scan of 467 F2 progeny of a broiler x layer cross was conducted to identify quantitative trait loci (QTL) affecting the rate of growth of the tail, wing and back feathers, and the width of the breast feather tract, at three weeks of age. 2. Correlations between the traits ranged from 0·36 to 0·61. Males had longer tail and wing feathers and shorter back feathers than females. Breast feather tract width was greater in females than males. 3. QTL effects were generally additive and accounted for 11 to 45% of sex average feather lengths of the breeds, and 100% of the breast feather tract width. Positive and negative alleles were inherited from both lines, whereas the layer allele was larger than the broiler allele after adjusting for body weight. 4. A total of 4 genome-significant and 4 suggestive QTL were detected. At three or 6 weeks of age, 5 of the QTL were located in similar regions as QTL for body weight. 5. Analysis of a model with body weight at three weeks as a covariate identified 5 genome significant and 6 suggestive QTL, of which only two were coincident with body weight QTL. One QTL for feather length at 148 cM on GGA1 was identified at a similar location in the unadjusted analysis. 6. The results suggest that the rate of feather growth is largely controlled by body weight QTL, and that QTL specific for feather growth also exist.

Feather growth rate and mass in Nearctic passerines with variable migratory behavior and molt pattern

ABSTRACT. Bird species vary greatly in the duration of their annual complete feather molt. However, such variation is not well documented in birds from many biogeographic areas, which restricts our understanding of the diversification of molt strategies. Recent research has revealed that molt duration can be estimated in passerines from ptilochronology-based measurements of the growth rate of their tail feathers. We used this approach to explore how molt duration varied in 98 Nearctic species that have different migratory strategies and molt patterns. As previously documented for Palearctic species, migration was associated with a shortening of molt duration among species that molted during summer on their breeding range. However, molts of winter-molting migratory species were as long as those of summer-molting sedentary species, which suggests that winter molt also allows Nearctic migrants to avoid the temporal constraints experienced during summer. Our results also suggest that migratory species that un...

The phenology of molting, breeding and their overlap in central Amazonian birds

The energetically challenging periods of molting and breeding are usually temporally separated in temperate birds, but can occur simultaneously in tropical birds, a condition known as molt–breeding overlap. Here, we document great variation in the timing and duration of molting and breeding, and in the extent of molt–breeding overlap, among 87 species of understory passerines in central Amazonia. We analyzed molt and breeding from 26 871 birds captured over a 30‐yr period near Manaus, Brazil. Although most species typically bred during the late dry season (about October through January), many thamnophilids apparently bred year‐round, whereas a few other species from a variety of families bred mainly during the wet season (about January through May). Of all breeding birds with an active brood patch, 12.7% were simultaneously molting. Molt–breeding overlap was more frequently observed among suboscines (13.3%), especially thamnophilids (23.0%), than oscines (6.4%). Some families had <5% molt–breeding overlap frequency, including Tyrannidae (4.4%), Tityridae (0.0%), Pipridae (1.5%), Turdidae (0.0%), and Thraupidae (0.0%), indicating that not all tropical species exhibit molt–breeding overlap. Among 31 well‐sampled species (n ≥15 brood patches), variation in molt–breeding overlap frequency was positively correlated with each species’ average duration of flight feather replacement (range 98–301 d). We also measured feather growth rates of individual birds in nine species; in five of these, slower‐growing feathers increased with an individual's probability of having molt–breeding overlap. Among furnariids, molt–breeding overlap occurred either at the beginning or end of the molt cycle, suggesting that physiological mechanisms typically separate molting from breeding. Thamnophilids showed a much different pattern; molt–breeding overlap occurred at any stage of feather replacement, apparently not regulated to be independent of breeding. These results reveal substantial life‐history variation among Amazonian birds. Future work to resolve the physiological regulation of molting and breeding in tropical birds will greatly contribute to understanding these patterns and their relevance to avian diversity.

Winter body condition in relation to age, sex and plumage ornamentation in a migratory songbird

Winter body condition may play important roles in the life history of migratory birds, but it is difficult to estimate. We used the growth rate of winter‐grown tail feathers of Collared Flycatchers Ficedula albicollis as an indicator of winter body condition, comparing this trait between age classes and sexes and relating it to plumage ornamentation (forehead and wing patch sizes). Adults and males were in better nutritional condition during winter, as indicated by their faster tail feather growth rates, than were yearlings and females, respectively, which could indicate differences in individual quality and foraging ability with age, or age‐ and sex‐related winter habitat segregation. However, feather growth rate was related neither to the size of the winter‐grown forehead patch nor to the size of the summer‐grown wing patch, suggesting weak condition‐dependence for the winter‐grown ornament and complex life‐history consequences for the summer‐grown ornament.

Developmental integration of feather growth and pigmentation and its implications for the evolution of diet‐derived coloration

Variation in avian coloration is produced by coordinated pigmentation of thousands of growing feathers that vary in shape and size. Although the functional consequences of avian coloration are frequently studied, little is known about its developmental basis, and, specifically, the rules that link feather growth to pigment uptake and synthesis. Here, we combine biochemical, modeling, and morphometric techniques to examine the developmental basis of feather pigmentation in house finches (Carpodacus mexicanus)--a species with extensive variation in both growth dynamics of ornamental feathers and their carotenoid pigmentation. We found that the rate of carotenoid uptake was constant across a wide range of feather sizes and shapes, and the relative pigmented area of feathers was independent of the total amount of deposited carotenoids. Analysis of the developmental linkage of feather growth and pigment uptake showed that the mechanisms behind partitioning the feather into pigmented and nonpigmented parts and the mechanisms regulating carotenoid uptake into growing feathers are partially independent. Carotenoid uptake strongly covaried with early elements of feather differentiation (the barb addition rate and diameter), whereas the pigmented area was most closely associated with the rate of feather growth. We suggest that strong effects of carotenoid uptake on genetically integrated mechanisms of feather growth and differentiation provide a likely route for genetic assimilation of diet-dependent coloration.

Carotenoid-based plumage coloration in golden-crowned kinglets Regulus satrapa: pigment characterization and relationships with migratory timing and condition

Carotenoid-based ornamental coloration has long been proposed to honestly signal quality due to its dependence on individual condition. Because migration can be one of the most stressful periods of an animal's annual cycle, developing colourful plumage may be particularly challenging for species in which migration and moult periods overlap or occur sequentially. The purpose of this study was to investigate pigmentary and condition-dependent bases of carotenoid colour variation in a small migratory passerine, the golden-crowned kinglet Regulus satrapa (Family Regulidae). We captured 186 male and female kinglets of various ages during fall migration in southwestern Ontario, Canada and recorded arrival date, body condition index, fat and pectoral muscle scores, wing mite infestation, and feather growth rate as measures of condition. We quantified crown coloration using reflectance spectrometry and analyzed feather carotenoids using high-performance liquid chromatography. Yellow crown feathers of female kinglets contained only yellow hydroxycarotenoids, whereas orange feathers of males harboured a suite of eight carotenoid pigments. Males with longer wavelength orange crown hues deposited greater concentrations of ketocarotenoids, especially canthaxanthin. Female kinglets with longer wavelength crown hues and males with longer wavelength crown hues and more saturated crown coloration left for migration earlier in the year. Females with longer wavelength crown hues had fewer feather mites and tended to be in better condition. However, male kinglets with more saturated coloration possessed smaller pectoral muscles. This is the first study to identify plumage carotenoids in this North American bird family and to determine the pigmentary basis for both inter- and intrasexual colour variation. Our results provide further support for the condition-dependence of carotenoid coloration and suggest that ornamental elaboration in both sexes may encode information about fall condition and migratory performance.

Using ptilochronology to determine daily mercury deposition in feathers of nestling waterbirds

Feathers are commonly used biomarkers of mercury (Hg) contamination in waterbird species. Most studies that analyze waterbird feathers for Hg content report concentrations on a per-unit mass basis. While this is appropriate for intraspecific comparisons, we suggest a more effective method for studies comparing multiple species of similar size and with similar foraging habits. Ptilochronology is a technique for determining the rate of feather growth in individuals based on their nutritional condition. When paired with Hg analysis, feather growth rates can be used to calculate the average daily Hg deposition into a feather. In this study we used this technique in comparison with the commonly used metric of Hg per-unit feather mass in two waterbird species. Average daily Hg deposition into feathers was 26.7% more sensitive to differences in Hg between the two species, suggesting that this may be a more biologically meaningful metric to use in interspecific comparisons.

How Much Variation in the Molt Duration of Passerines can be Explained by the Growth Rate of Tail Feathers?

ABSTRACT. In birds, molt duration is an important trait that can affect plumage functionality and, consequently, the fitness of individuals. However, knowledge about the factors that affect variation in molt speed is sparse, mostly because of the methodological difficulties of studying avian molt. We used a ptilochronology-based approach to estimate the rate at which tail feathers were produced during molt to shed light on the relationship between molt duration and feather growth rate. For that purpose, we used three data sets. First, we tested whether the average molt durations of 22 passerine species were correlated with the mean growth rates of their feathers, using both conventional and phylogenetically corrected statistical procedures. Second, we explored this same association among captive Great Tits (Parus major). And third, we took advantage of the biannual complete molt of Willow Warblers (Phylloscopus trochilus) to examine whether the feathers synthesized during their short summer molt grew fast...

Sex‐dependent response of primary moult to simulated time constraints in the rock sparrow Petronia petronia

There is growing evidence that moult speed affects plumage quality. In many bird species, males and females differ in terms of breeding effort, survival expectation and the relationship between fitness and plumage quality. Consequently, differences in moult strategies between the sexes can be expected. The aim of this study was to assess whether, under simulated time constraints and with no parental investment in the previous breeding season, males and females differed in: a) timing and duration of primary moult, b) growth rates of individual primary feathers, and c) number of concurrently growing feathers. We investigated the effect of time constraints generated by a treatment consisting of two decreasing photoperiods (slow changing photoperiod, SCP=2 min day−1 and fast changing photoperiod, FCP=8 min day−1) on the primary post‐nuptial moult of captive rock sparrows Petronia petronia. Females started to moult on average 14 and 15 days later than males in both experimental groups. Primary moult duration was 10 (FCP) and 24 (SCP) days longer in males than in females, and, within sex, 34 (females) and 48 (males) days longer in SCP birds than in FCP ones. Females renewed a larger number of primaries simultaneously (5.7% in FCP and 12.8% in SCP) and had a higher total daily feather mass grown (9.9% in FCP and 22.4% in SCP), even though daily growth rates of individual primaries did not differ between sexes. As a result, males and females completed their primary moult at the same time within treatment. The observed differences in timing, duration and energy allocation for primary moult between the sexes probably have a genetic basis, as birds did not engage in reproduction during the preceding breeding season.

Fitness consequences of timing of breeding in birds: date effects in the course of a reproductive episode

In seasonal environments, avian reproductive performance almost generally declines in the course of the season. Quantifying the associated fitness consequences of timing of breeding, i.e. of date‐related factors, is important for understanding the evolution of temporal patterns in avian life‐histories and for predicting consequences of climate change. The seasonal decline can also be caused by an effect of parental quality: individuals with high phenotypic quality may breed early. The results of existing experimental studies investigating whether date or quality effects cause the seasonal decline are inconsistent, indicating that both mechanisms might be involved. However, it remains unclear to what extent the confounding effect of quality occurs and what the fitness consequences of timing per se over a whole breeding episode are. In a cross‐fostering experiment using the barn swallows’ second broods we evaluated the causes for the seasonal decline in reproductive performance for three distinct periods of a reproductive attempt, the early nestling period, the late nestling period and the post‐fledging period, and we assessed the overall fitness consequences of timing per se. A seasonal decline in juvenile feather growth rate was mainly due to date effects in the late nestling period, although we determined quality effects during early nestling development. Date effects on survival were present in the post‐fledging period, but not in the nestling period. The decline in feather length due to date effects in the nestling period accounted for 9% of the seasonal decline in post‐fledging survival, whereas date effects arising only in the post‐fledging period caused 91% of the decline. These results suggest that date effects increase in the course of a reproductive episode. Thus, the benefits of an early timing of breeding can be quantified only when considering also the post‐fledging period. We suggest that the timing of breeding evolved through a trade‐off between date‐related benefits and quality‐related costs of early breeding.

Genetic and environmental effects on a condition‐dependent trait: feather growth in Siberian jays

Condition, defined as the amount of 'internal resources' an individual can freely allocate, is often assumed to be environmentally determined and to reflect an individual's health and nutritional status. However, an additive genetic component of condition is possible if it 'captures' the genetic variance of many underlying traits as many fitness-related traits appear to do. Yet, the heritability of condition can be low if selection has eroded much of its additive genetic variance, or if the environmental influences are strong. Here, we tested whether feather growth rate - presumably a condition-dependent trait - has a heritable component, and whether variation in feather growth rate is related to variation in fitness. To this end, we utilized data from a long-term population study of Siberian jays (Perisoreus infaustus), and found that feather growth rate, measured as the width of feather growth bars (GB), differed between age-classes and sexes, but was only weakly related to variation in fitness as measured by annual and life-time reproductive success. As revealed by animal model analyses, GB width was significantly heritable (h(2) = 0.10 +/- 0.05), showing that this measure of condition is not solely environmentally determined, but reflects at least partly inherited genetic differences among individuals. Consequently, variation in feather growth rates as assessed with ptilochronological methods can provide information about heritable genetic differences in condition.

Allometry of the Duration of Flight Feather Molt in Birds

Author SummaryThe pace of life varies with body size and is generally slower among larger organisms. Larger size creates opportunities but also establishes constraints on time-dependent processes. Flying birds depend on large wing feathers that deteriorate over time and must be replaced through molting. The lengths of flight feathers increase as the 1/3 power of body mass, as one expects for a length-to-volume ratio. However, feather growth rate increases as only the 1/6 power of body mass, possibly because a two-dimensional feather is produced by a one-dimensional growing region. The longer time required to grow a longer feather constrains the way in which birds molt, because partially grown feathers reduce flight efficiency. Small birds quickly replace their flight feathers, often growing several feathers at a time in each wing. Larger species either prolong molt over two or more years, adopt complex patterns of multiple feather replacement to minimize gaps in the flight surface, or, among species that do not rely on flight for feeding, simultaneously molt all their flight feathers. We speculate that the extinct 70-kg raptor, Argentavis magnificens, must have undergone such a simultaneous molt, living off fat reserves for the duration.

Repeatable Length and Mass but Not Growth Rate of Individual Feathers between Moults in a Passerine Bird

Plumage is regularly replaced over the lifetime of birds, and, consequently, variable conditions experienced during plumage development can affect feather growth rate or the expression of feather characteristics. We assessed intra-individual constancy in the length, mass, and growth rate of tail feathers between complete moults of 13 free-liv- ing individual Blackcaps Sylvia atricapilla. Between-moult repeatability was high for the length and mass, but not for growth rate of individual tail feathers. The feathers produced in the later moult grew at a slower rate, which shows that the time to produce similar feathers increases with age.

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla. As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.

Case Study: A Split–brood Comparison of Formula for Nestling Songbirds (FoNS ) versus Three Facility–specific Diets

Thirty–four nestling birds of seven species were used to test whether Formula for Nestling Songbirds (FoNS ) would result in better weight gain, feather growth, and survival rates compared to inexpensive control diets. Ten broods, of 2 to 5 birds, were split into two groups. Some of each brood received FoNS while others received one of three species specific and facility–formulated control diets. Four species (Bewick’s wren [Thryomanes bewickii], red–eyed vireo [Vireo olivaceus], violet–green swallow [Tachycineta thalassina], and black–capped chickadee [Poecile atricapillus]), received a control diet consisting of dog food supplemented with bone meal, brewer’s yeast, and protein powder. With the exception of the swallows, birds on this control diet generally lost weight, and three of the five died; whereas all six birds from the same broods survived and gained weight on FoNS . Two species, (house sparrow [Passer domesticus] and house finch [Carpodacus mexicanus]) received control diets of chick starter with cat food, hard boiled eggs, bone meal powder, and peanut butter, and one species (European starling [Sturnus vulgaris]) received a control diet based on dog food. For these species, the FoNS diet and control diets showed similar development, growth, and no deaths. The authors suggest that FoNS diet is advantageous, especially for small, insectivorous species, but some inexpensive alternatives may be equally suitable for certain species.

INTRAFEATHER AND INTRAINDIVIDUAL VARIATION IN THE STABLE-HYDROGEN ISOTOPE (δD) CONTENT OF RAPTOR FEATHERS

Abstract Stable-hydrogen isotope ratios (deuterium:protium; δD) in feathers enable researchers to evaluate patterns of avian movement and to estimate the source areas of migratory birds. However, variation in feather δD remains inadequately described, thus confounding inferences of avian movement and origin. We assessed variation within a feather and among feathers within and between tracts in three species of immature raptors. Within contour feathers, measurements of δD increased from a distal section to an adjacent, proximal section; the magnitude of δD increase varied with raptor species. Furthermore, contour and flight feathers differed systematically in their δD content. Two explanations for intrafeather and intraindividual variation warrant further investigation: (1) hydrogen isotope fractionation associated with feather growth rate, and (2) the incorporation of temporal variation in environmental δD into growing feathers. We consider these explanations for raptors and passerines, which seemingly di...

Differential Development of Breeding Plumage in Adult and Second-Year Male RuffsPhilomachus pugnax

We studied the development of breeding plumage in 641 Ruff Philomachus pugnax males during spring in 2004–2006 in southern Belarus with special emphasis on the growth rate of ruff and tuft feathers, which are the most colourful and spectacular parts of the male plumage. During northward migration, male Ruffs arriving in the end of March or early April showed a mixture of winter and striped feathers. The first adults with fully developed breeding plumage were recorded in the last 10 days of April and the proportion of individuals with breeding plumage gradually increased through spring. However, individuals with at least traces of winter or striped plumage were caught during the whole study period. Birds did not shed the whole winter plumage at once, which is typical for Ruff males in western Europe. Immature (second year) birds were less advanced in developing breeding plumage than adults. Only 20% of all 55 trapped second-year males were in full breeding plumage. During the main migration period there were no annual differences in the growth rate of breeding plumage feathers. Ruff lengths increased 1.6 mm/day in adults and 1.5 mm/day in second-year males, whereas the tuft grew 1.1 mm/day in adults and 1.0 mm/day in immatures. Although growing rates of ruff and tuft feathers were similar in immatures and adults, immature males started to moult into breeding plumage later than adults, resulting in less developed breeding plumage when they departed to the breeding grounds. As second-year males had lower body mass and slower body mass increase than adults, it was proposed that energetic constraints during migration are the main reason of delayed moult into breeding plumage in immature birds.

Feather development under environmental stress: lead exposure effects on growth patterns in Great Tits Parus major

Capsule Regrowth rate of tail feathers is more strongly affected compared to feather length and symmetry. Aims To assess the value of avian feathers as bioindicators. Methods The origin and persistence of fluctuating asymmetry (FA) in homologous pairs of regrowing feathers was studied in captive birds under different levels of environmental stress, respresented by exposure to lead (Pb). Homologous feathers of individually housed birds were plucked synchronously or with a delay of seven days. We measured growth rate, regeneration time, final size and FA of regrown feathers and related them to Pb stress. Results Asymmetry decreased as feathers reached their final length. This was not due to compensatory growth but rather a consequence of the programmed growth trajectory of single feathers. Tail feathers grown under higher Pb pollution showed increased regeneration times, decreased growth rates and shorter lengths, but no changes in development times nor in FA. For differences between both (i) original and induced feathers and (ii) control and Pb treatment, effect sizes of parameters related to feather development (growth bar width, growth rate, regeneration and development time) were consistently larger than those related to the resulting phenotype (feather length and FA). Conclusions Growth bar widths in particular provide an applicable, sensitive and reliable indicator of adverse conditions such as Pb pollution and experimental conditions. In general, phenotypic characteristics that retrospectively allow estimation of growth rates may be more suitable for monitoring environmental stress than sizes or asymmetries of full-grown traits.

Parasite‐mediated growth patterns and nutritional constraints in a cavity‐nesting bird

1. Trade-offs between growth and immunity of nestling birds can be influenced by parasites, but the magnitude of these effects may depend on availability of critical dietary nutrients. Owing to their importance for both immune system function and growth, dietary carotenoids have the potential to mediate parasite-induced developmental strategies of avian hosts. 2. The effects of ectoparasitic blow flies Protocalliphora spp. and dietary carotenoids (lutein and zeaxanthin) on immune function and patterns of growth in nestling mountain bluebirds Sialia currucoides were investigated by combining parasite removal and carotenoid supplementation treatments in a 2 x 2 design. 3. Supplemental carotenoids enhanced nestlings' T-cell-mediated immune response following intradermal injection of phytohaemagglutinin. 4. The effect of carotenoid supplementation on rate of mass gain depended on whether broods were exposed to parasites: among parasitized broods, those receiving supplemental carotenoids gained mass more rapidly than nonsupplemented broods, whereas there was no effect of supplemental carotenoids on growth of mass in broods that had parasites removed. This suggests that additional dietary carotenoids allowed nestlings to compensate for the otherwise detrimental effects of parasites on mass gain. For length of the eighth primary feather at fledging, early and late broods differed in their response to parasitism: early broods showed an increase in feather length when parasites were removed, while nestlings in late broods had shorter feathers in the absence of parasites. We suggest that this may reflect within-season variation in parasite-mediated growth strategies of nestlings. 5. Maternal condition was positively associated with mass, condition and rate of feather growth of offspring under all conditions, and also influenced nestling immunocompetence, but only in the absence of parasites. 6. We conclude that dietary carotenoids alleviate some of the detrimental effects of parasites on nestling birds; however, parasites also appear to specifically influence other growth and resource allocation strategies, and possibly constrain maternal or genetic effects on offspring phenotype, irrespective of dietary carotenoid availability.