Prenuptial Molt Research Articles

Plumage dimorphism and moult patterns of weaverbirds – a review

The colourful plumages of weaverbirds (family Ploceidae, with 118 species recognised) provide a well-known and in part iconic showcase of sexual selection, with widespread sexual dimorphism (93 species) as well as seasonal dimorphism (62 species). The tremendous diversity in both the type and degree of plumage dimorphism is poorly known, which calls for a better and extended understanding of the highly variable moult patterns in this group. Here, we lay a foundation for this by reviewing and discussing what is known of the timings and patterns of moult in the Ploceidae. The extent of post-juvenile moult is poorly known in weavers: it is known to be complete in 10 species, partial in two species, and either complete or partial in six species. However, it is likely that there is more variation than previously thought. Weavers typically have a complete moult after breeding, and before breeding a partial prenuptial moult, restoring the male (and, in some species, female) breeding plumage; this usually involves only body feathering, one exception being the Seychelles Fody Foudia sechellarum, which has a complete prenuptial moult. We compiled primary moult data based on the proportion of feather mass grown, for 11 weaver species, three of which are newly reported on. Timing of primary moult is discussed in relation to sex, location and annual timing. The duration of primary moult varies widely between and within species, from 46 days (1.5 months) in a widowbird to 215 days (7 months) in an arid species. Weavers in arid regions have longer moult durations than weavers in other regions. Examples of the overlap of primary moult and breeding are presented.

Seasonal but not sex-biased gene expression of the carotenoid ketolase, CYP2J19, in the sexually dichromatic southern red bishop (Euplectes orix).

Intense red colours in birds are often owing to ketocarotenoids (KCs). In many land birds, KCs are oxidized from dietary yellow precursors, presumably by the avian carotenoid ketolase CYP2J19, the regulation and constraints of which have important implications for condition-dependence and honest signalling of carotenoid colour displays. We investigated hepatic CYP2J19 gene expression in the seasonally and sexually dichromatic southern red bishop (Euplectes orix) in relation to season, sex, progression of the prenuptial moult, testis size, body condition, redness and circulating sex steroids. A coloration function of CYP2J19 is supported by a seasonal upregulation prior to and during the carotenoid-depositing stage of the male prenuptial moult. However, CYP2J19 expression was similarly high in females (which do not moult prenuptially), and remained high in males after moult, suggesting additional or alternative roles of hepatic CYP2J19 or its products, such as detoxification or antioxidant functions. In males, the CYP2J19 upregulation preceded and was unrelated to the rise in plasma testosterone, but was correlated with androstenedione, probably of adrenal origin and compatible with luteinizing hormone-induced and (in females) oestrogen-suppressed moult. Finally, contrary to ideas that carotenoid ketolation rate mediates honest signalling of male quality, CYP2J19 expression was not related to plumage redness or male body condition.

The extent of white plumage patches in female Pied Flycatchers Ficedula hypoleuca is negatively associated with corticosterone concentration in partly unpigmented feathers

Sexually and socially selected signals are predicated to express the present and past condition of individuals and thereby their capacity to cope with environmental challenges. The concentration of corticosterone in feathers (CORTf) has been validated as a marker of activation of the hypothalamic-pituitary axis in birds during feather development. Measurements of CORTf can thus express the physiological stress of migratory birds during non-breeding periods of the annual cycle when feathers moulted in the wintering range are analysed. Thus, negative trends of ornament expression with CORTf may be predicted. Unpigmented plumage patches may constitute signals of phenotypic quality and are used in many species in social interactions by both sexes. Pied Flycatchers of both sexes Ficedula hypoleuca show white forehead and wing patches during the breeding season, of which the forehead and part of the wing patch (tertials) are moulted on the wintering grounds before spring migration. These patches are used as signals in social and sexual interactions and their expression has been previously related to several indicators of physiological status. We collected tertials of females during nestling provisioning in a Spanish population and determined their CORTf in the laboratory. CORTf was thus measured in some of the prenuptially moulted feathers used as signals of social dominance by breeding females. According to AICc model selection, CORTf turned up in all the best models explaining forehead patch size and in most best models for wing patch size when controlling for age and size. Both forehead and wing patch sizes were negatively related to CORTf, although the trend for wing patch size was not significant. This indicates that breeding females expressing large unpigmented patches experienced low levels of stress during the prenuptial moult in their winter quarters, or that strongly signalling individuals present a lower level of activation of the HPA axis.

Phenotypic flexibility of glucocorticoid signaling in skeletal muscles of a songbird preparing to migrate

Glucocorticoids are commonly associated with responses to stress, but other important functions include homeostatic regulation, energy metabolism and tissue remodeling. At low circulating levels, glucocorticoids bind to high-affinity mineralocorticoid receptors (MR) to activate tissue repair and homeostasis (anabolic pathways), whereas at elevated levels, glucocorticoids bind to glucocorticoid receptors (GR) to activate catabolic pathways. Long distance migrations, such as those performed by Gambel's white-crowned sparrows (Zonotrichia leucophrys gambelii), require modification of anatomy, physiology and behavior. Plasma corticosterone (CORT) increases in association with impending departure and flight and may promote muscle-specific anabolic states. To test this idea, we explored glucocorticoid signaling in the pectoralis (flight) and gastrocnemius (leg) muscles of male sparrows on the wintering grounds at three stages leading up to spring departure: winter (February), pre-nuptial molt (March), and pre-departure (April). CORT was detected in plasma and in both muscles, but measures of CORT signaling differed across muscles and stages. Expression of 11β-hydroxysteroid dehydrogenase (11β-HSD) Type 2 (inactivates CORT) increased in the pectoralis at pre-departure, whereas 11β-HSD Type 1 (regenerates CORT) did not change. Neither of the two 11β-HSD isoforms was detectable in the gastrocnemius. Expression of MR, but not GR, was elevated in the pectoralis at pre-departure, while only GR expression was elevated at pre-nuptial molt in gastrocnemius. These data suggest that anabolic functions predominate in the pectoralis only while catabolic activity is undetected in either muscle at pre-departure.

Preparing to migrate: expression of androgen signaling molecules and insulin-like growth factor-1 in skeletal muscles of Gambel's white-crowned sparrows.

Migratory birds, including Gambel's white-crowned sparrows (Zonotrichia leucophrys gambelii), exhibit profound modifications of skeletal muscles prior to migration, notably hypertrophy of the pectoralis muscle required for powered flight. Muscle growth may be influenced by anabolic effects of androgens; however, prior to spring departure, circulating androgens are low in sparrows. A seasonal increase in local androgen signaling may occur within muscle to promote remodeling. We measured morphological parameters, plasma and tissue levels of testosterone, as well as mRNA expression levels of androgen receptor, 5α-reductase (converts testosterone to 5α-dihydrotestosterone), and the androgen-dependent myotrophic factor insulin-like growth factor-1. We studied the pectoralis muscle as well as the gastrocnemius (leg) muscle of male sparrows across three stages on the wintering grounds: winter (February), pre-nuptial molt (March), and pre-departure (April). Testosterone levels were low, but detectable, in plasma and muscles at all three stages. Androgen receptor mRNA and 5α-reductase Type 1 mRNA increased at pre-departure, but did so in both muscles. Notably, mRNA levels of insulin-like growth factor-1, an androgen-dependent gene critical for muscle remodeling, increased at pre-departure in the pectoralis but decreased in the gastrocnemius. Taken together, these data suggest a site-specific molecular basis for muscle remodeling that may serve to enable long-distance flight.

Effects of thyroid hormone manipulation on pre-nuptial molt, luteinizing hormone and testicular growth in male white-crowned sparrows (Zonotrichia leuchophrys gambelii)

Most seasonal species rely on the annual change in day length as the primary cue to appropriately time major spring events such as pre-nuptial molt and breeding. Thyroid hormones are thought to be involved in the regulation of both of these spring life history stages. Here we investigated the effects of chemical inhibition of thyroid hormone production using methimazole, subsequently coupled with either triiodothyronine (T3) or thyroxine (T4) replacement, on the photostimulation of pre-nuptial molt and breeding in Gambel’s white-crowned sparrows (Zonotrichia leuchophrys gambelii). Suppression of thyroid hormones completely prevented pre-nuptial molt, while both T3 and T4 treatment restored normal patterns of molt in thyroid hormone-suppressed birds. Testicular recrudescence was blocked by methimazole, and restored by T4 but not T3, in contrast to previous findings demonstrating central action of T3 in the photostimulation of breeding. Methimazole and replacement treatments elevated plasma luteinizing hormone levels compared to controls. These data are partially consistent with existing theories on the role of thyroid hormones in the photostimulation of breeding, while highlighting the possibility of additional feedback pathways. Thus we suggest that regulation of the hypothalamic pituitary gonad axis that controls breeding may be more complex than previously considered.

Subspecies-specific features of molt in the common chiffchaff (Phylloscopus collybita L.) from Europe and Western Siberia

The patterns of molt in the chiffchaffs of the nominotypical subspecies from central Europe and eastern European subspecies from the Ladoga region are compared with that of the Siberian subspecies from the lower Ob region. The onset dates of postjuvenile molt in the Siberian subspecies are controlled endogenously, while in the nominative and, probably, eastern European subspecies they are under photoperiodic control. The molt of chiffchaffs from central Europe is no less complete than in birds from the Ladoga region and more complete than in Siberian birds. The individual duration of molt is the shortest in Siberian birds and the longest in central European birds, being intermediate in birds from the Ladoga region. Differences in the pattern of postnuptial molt between chiffchaffs from the Ladoga and Ob regions concern its duration and completeness. The completeness of molt proved to be reduced in half of examined birds from the Ladoga region and in some birds from the Ob region. The duration of complete molt in birds from the Ob region is smaller (40 days vs. 60 days in birds from the Ladoga region) at a longer daylight period, which is evidence for a higher rate of feather growth. The completeness and timing of prenuptial molt do not differ between the subspecies.

Early-moulting Red-backed Fairywren males acquire ornamented plumage in the absence of elevated androgens

ABSTRACTSexual ornaments, including plumage ornamentation, are often studied during breeding periods even though signal development can take place months earlier. This temporal disconnect potentially obscures the proximate mechanisms that underlie signal expression and development. We studied the correlation between androgen levels and expression of ornamented plumage in adult Red-backed Fairywrens (Malurus melanocephalus) 4–6 months before breeding, when signal production occurs in some, usually older, males. We found that, during this period, ornamented males, unornamented males and females all had low plasma androgen levels that did not differ from each other. Variation in androgen levels was unrelated to phenotype or moult. These findings differ from previous research conducted immediately prior to breeding in a different population of this species, which used correlative and experimental work to demonstrate that testosterone induces prenuptial moult and acquisition of ornamented plumage in younger males. Our study demonstrates that mechanisms contributing to signal production may vary within and among populations in relation to temporal, age-dependent, or geographic parameters. These results highlight the complexity of hormonal pathways to signal production, and the importance of studying signal acquisition throughout the entire period when signals are produced, as studies conducted at different time points may have quantitatively different results.

Behavioral and physiological traits of migrant and resident white-crowned sparrows: a common garden approach.

To accommodate a migratory life history, migrants express a greater number of physiological and behavioral stages per annum than residents and are thus considered to have higher finite state diversity (FSD). To investigate the physiological mechanisms and constraints associated with migration, direct comparison of two subspecies of white-crowned sparrow - migrant, Zonotrichia leucophrys gambelii, and resident, Z. l. nuttalli - were made under common garden conditions of photoperiod and housing, as birds progressed from winter through the vernal life history stages. We tested the hypothesis that migrants (higher FSD) respond differently than residents (lower FSD) to the initial predictive cue, photoperiod, to initiate and integrate the progression of vernal stages of prenuptial molt, migration and development of breeding. If differences in vernal phenology were noted, then the basis for the distinctions was considered genetic. Results indicate that (1) residents had a lower threshold to vernal photoperiod with elevations of plasma androgen, growth and development of reproductive structures preceding those of migrants; (2) only migrants displayed prenuptial molt, preparations for migration and migratory restlessness; and (3) neither baseline nor stress-induced plasma corticosterone differed across subspecies, suggesting energetic demands of the common garden were insufficient to induce a differential adrenocortical response in either subspecies, highlighting the impact of environmental conditions on corticosterone secretion. Thus, in a common garden experiment, Z. l. gambelii responds differently to the initial predictive cue, photoperiod, to initiate and execute the vernal stages of molt, migration and development of breeding in comparison to the shared stage of breeding with Z. l. nuttalli, confirming a genetic basis for the subspecies differences.

Testosterone activates sexual dimorphism including male-typical carotenoid but not melanin plumage pigmentation in a female bird.

In males it is frequently testosterone (T) that activates the expression of sexually selected morphological and behavioral displays, but the role of T in regulating similar traits in females is less clear. Here, we combine correlational data with results from T and gonadotropin-releasing hormone (GnRH) manipulations in both sexes to assess the role of T in mediating sexually dimorphic coloration and morphology in the red-backed fairy-wren (Malurus melanocephalus). We show that: (1) natural variation in female expression of ornamental traits (darkened bills and red back feathers) is positively associated with age and circulating androgen titres, (2) females have the capacity to express most male-typical traits in response to exogenous T, including carotenoid-pigmented body plumage, shorter feathers, darkened bill and enlarged cloacal protuberance, but (3) appear constrained in production of male-typical melanin-pigmented plumage, and (4) low androgen levels during the pre-nuptial molt, probably because of low ovarian capacity for steroid production (or luteinizing hormone sensitivity), prevent females from developing male-like ornamentation. Thus, females appear to retain molecular mechanisms for hormonally regulated male-typical ornamentation, although these are rarely activated because of insufficient production of the hormonal signal.

Feather and faecal corticosterone concentrations predict future reproductive decisions in harlequin ducks (Histrionicus histrionicus).

Understanding sources of reproductive variation can inform management and conservation decisions, population ecology and life-history theory. Annual reproductive variation can drive population growth rate and can be influenced by factors from across the annual cycle (known as carry-over effects). The majority of studies, however, focus solely on the role of current environmental events. Past events often influence future reproductive decisions and success but can be logistically difficult to collect and quantify, especially in migratory species. Recent work indicates that glucocorticoids may prove good indicators to evaluate carry-over effects across life-history transitions. Here, we evaluated three different measures of glucocorticoid physiology (feathers, faeces and plasma) to evaluate the predictability of future breeding decision in the harlequin duck (Histrionicus histrionicus). We collected tail and back feathers, plasma and faeces for glucocorticoid analysis, and fitted female harlequin ducks with very high-frequency transmitters to track their breeding decisions. Both back feathers (moulted immediately before the current season) and faecal glucocorticoid metabolites were identified as important predictive factors of reproductive decisions; high concentrations of glucocorticoid metabolites in back feathers and faeces predicted a higher likelihood of reproductive deferral for the year. Although back and tail feather corticosterone concentrations were correlated, tail feathers (moulted at the end of the previous breeding season) did not predict breeding decisions. Plasma corticosterone concentrations were collected over too broad a time range after capture to be useful in this study. This study demonstrates the utility of non-invasive corticosterone metrics in predicting breeding decisions and supports the use of feathers to measure carry-over effects in migratory birds. With this technique, we identified the prenuptial moult as an important life-history phase that contributes to reproductive decisions. Identification of critical life-history phases is paramount to efficient management of species.

Regulatory mechanisms for the development of the migratory phenotype: Roles for photoperiod and the gonad

This article is part of a Special Issue “Energy Balance”.Male white-crowned sparrows, Zonotrichia leucophrys gambelii, were studied to investigate roles of natural day length and the testes in regulating development and expression of the vernal migration phenotype. Previous work suggested that a pulse of androgen during winter months followed by the vernal increase in photoperiod promotes fueling (fat deposition) to support long distance flight; however, other traits required for successful migration remain untested. To investigate these points, birds were captured on their wintering grounds and castrated prior to winter solstice following Mattocks (1976). A subset of the castrates received 8mm Silastic implants of testosterone (T-castrates) and others blank implants (Blank-castrates) for 16days in February. Shams were surgical controls. Migratory traits measured were as follows: 24h locomotor activity, prenuptial molt, body mass, fat score, flight muscle profile, cloacal protuberance (CPL) and plasma androgens measured over 28weeks divided into 3 experimental periods (pre-implant, implant, and post-implant). Under short day lengths, castration increased diurnal locomotor activity over Shams. Testosterone implants temporarily enhanced CPL, plasma androgens and flight muscle enlargement, but failed to induce migratory restlessness. Whereas all groups exhibited seasonal increases in mass, fat score and muscle profile, only Shams showed timely onset and completion of prenuptial molt and migratory restlessness. Thus, for castrated males exposed to naturally increasing day lengths, the organizational effects of a transient testosterone surge were not sufficient to actuate a timely spring molt and migratory behavior. A fully functional testis that can organize central processes is required for the entire expression of the spring migratory phenotype.

Photoperiodic regulation of seasonal reproduction, molt and body weight in the migratory male yellow-breasted bunting (Emberiza aureola)

Photoperiod has been shown to be a major source of temporal information regulating reproduction and associated functions in a number of avian species. We studied seasonal cycles of testicular volume, molt and body weight in natural and temperature-controlled conditions and under different artificial photoperiods in the yellow-breasted buntings. Buntings posses seasonal cycles of testicular volume, molt, body weight and fattening with no major difference between natural and temperature-controlled conditions. These cycles follow an annual solar cycle suggesting the possibility of their photoperiodic control. To confirm this, photosensitive birds were studied under 9L/15D (close to shortest day length), 12L/12D (equinox day length) and 14L/10D (close to longest day length) for 18 months. Buntings showed testicular growth followed by regression and development of photorefractoriness; molt and body weight change only under 12L/12D and 14L/10D but not under 9L/15D. Reinitiation of above responses did not occur following initial cycles under stimulatory photoperiods precluding the possibility of circannual rhythm involvement. Birds exhibited an incomplete prenuptial molt of body feathers during gonadal stimulation under long days followed by complete postnuptial molt of body and primary feathers that progressed with gonadal regression. Exposure of photosensitive birds to light–dark cycles constituting 9–16h of light/day suggested that daily photoperiod of about 12h or more is essential in inducing testicular growth and function. These results clearly indicate that buntings are capable of fine discrimination of photoperiodic information and use annual changes in day length as an environmental factor to time their seasonal responses.

Body condition influences sexual signal expression independent of circulating androgens in male red-backed fairy-wrens

Androgens play a major role in the regulation of sexual signal expression of male vertebrates. In this study we assessed the prevalent, yet largely untested, assumption that signal honesty is maintained through condition-dependent androgen regulation by experimentally manipulating body condition of male red-backed fairy-wrens (Malurus melanocephalus) through trimming several flight feathers before the prenuptial molt. In their first reproductive season males of this species exhibit androgen-regulated plasticity in plumage coloration, ranging from red/black (high androgens) to brown (low androgens). Red/black plumage is preferred by females and might be constrained by a negative relationship between body condition and androgen levels. We also evaluated whether corticosterone changes to altered conditional state mediate the relationship between condition and androgens. While we predicted that males with trimmed feathers would expend greater energy and thus be in poorer condition at the time of molt, they were counter-intuitively in better condition compared to control birds, likely as a consequence of subtle behavioral changes. These birds in better condition molted a greater proportion of red/black plumage, as predicted, and also molted more heavily. However, experimental and control birds did not differ in their androgen or corticosterone concentrations. Furthermore, analysis of long-term data from the same population revealed no correlation between condition and androgen levels. Collectively, these results challenge the notion that condition-dependent androgen regulation alone is responsible for maintaining the honesty of sexual signals and highlights the necessity of considering alternate explanations.

Male mate choice and male–male competition in the hermit crab Pagurus nigrofascia: importance of female quality

Male mate choice has recently been reported in some animals with male–male competition. In the laboratory, we examined whether males choose their mates based on female quality that was indicated by body size and/or days to prenuptial molt, and the effects of female quality on male–male competition in the hermit crab Pagurus nigrofascia. We collected samples from April to May 2009 at an intertidal shore in Hokkaido, Japan (41°N, 140°E). When a male simultaneously encountered two receptive females in the mate choice experiment, males chose females which require less time to molt. When a male guarding a female with less time to molt was challenged by an intruder, the guarding male defended the female for a longer period and was more likely to win the contest. These results indicate that male P. nigrofascia use time to molt to discriminate between females.

Sexually Selected Male Plumage Color Is Testosterone Dependent in a Tropical Passerine Bird, the Red-Backed Fairy-Wren (Malurus melanocephalus)

BackgroundSexual signals, such as bright plumage coloration in passerine birds, reflect individual quality, and testosterone (T) may play a critical role in maintaining signal honesty. Manipulations of T during molt have yielded mixed effects on passerine plumage color, in most cases delaying molt or leading to production of drab plumage. However, the majority of these studies have been conducted on species that undergo a post-nuptial molt when T is low; the role of T in species that acquire breeding plumage during a pre-nuptial molt remains largely unexplored.Methodology/Principal FindingsWe experimentally tested the effects of increased T on plumage color in second-year male red-backed fairy-wrens (Malurus melanocephalus), a species in which after-second-year males undergo a pre-nuptial molt into red/black (carotenoid and melanin-based) plumage and second-year males either assume red/black or brown breeding plumage. T treatment stimulated a rapid and early onset pre-nuptial molt and resulted in red/black plumage acquisition, bill darkening, and growth of the sperm storage organ, but had no effect on body condition or corticosterone concentrations. Control males molted later and assumed brown plumage. T treated males produced feathers with similar but not identical reflectance parameters to those of unmanipulated after-second-year red/black males; while reflectance spectra of red back and black crown feathers were similar, black breast feathers differed in UV chroma, hue and brightness, indicating a potentially age and plumage patch-dependent response to T for melanin- vs. carotenoid-pigmentation.Conclusions/SignificanceWe show that testosterone is the primary mechanism functioning during the pre-nuptial molt to regulate intrasexually variable plumage color and breeding phenotype in male red-backed fairy-wrens. Our results suggest that the effects of T on plumage coloration may vary with timing of molt (pre- vs. post-nuptial), and that the role of T in mediating plumage signal production may differ across age classes, plumage patches, and between pigment-types.

Role of gonadal and adrenal steroids and thyroid hormones in the regulation of molting in domestic goose

Plasma levels of testosterone (T), 17-β-estradiol (E2), progesterone (P4), dehydroepiandrosterone (DHEA), corticosterone (B), thyroxine (T4) and triiodothyronine (T3) were monitored during postnuptial and the prenuptial molt in domestic goose (Anser anser domesticus) in both sexes. 1. At the beginning of postnuptial molt (when the old, worn dawny-, and cover feathers' loss starts) in ganders, the levels of T, E2, P4 decrease while DHEA and B significantly increase. The elevated levels of T4 and low T3 concentrations characteristic of the last phase of the reproduction, remain unchanged. In layers, similar changes were observed, however, B decreases. 2. In the early phase of outgrowth of wing and cover feathers, plasma levels of T, E2 and P4 are low. Elevated B, DHEA and T4 concentrations decrease in ganders, while in layers DHEA increases and B and T4 levels remain unchanged. T3 increases in both sexes. 3. The subsequent intensive outgrowth period of wing- and cover feathers both in ganders and in layers is characterized by very low levels of T, E2, DHEA and T4, but P4 increased, and T3 concentration remain high. 4. At the end of postnuptial molt - when the outgrowth of dawny, cover-, and wing feathers stops - very low T, E2, P4, DHEA and T4 levels and and high T3 plasma levels were found in both sexes. Fast increase of plasma B was detected in ganders, while in geese, B concentration remain high. 5. During prenuptial molting (outgrowth of contour and tail feathers) low E2, P4 and T4, increasing T and DHEA, but very high T3 and B plasma concentration were measured in ganders. In layers, very low T, E2, P4, DHEA and T4 levels, and very high B and T3 levels were found. 6. At the beginning of the fall-winter sexual repose (postmolting stage) T, E2, P4, DHEA and T4 levels increase, T3 and B declines in both sexes. 7. In the subsequent phase of fall-winter period (preparatory stage) there is a further increase in T, P4 and T4, a fast increase of B and a decrease of E2, DHEA and T3 in ganders. In layers, T, P4 and DHEA decrease, B increases and the T4 and T3 do not change. 8. At the beginning of reproduction high T level, unchanged DHEA, slightly declined P4, and decreased E2, T4, T3 and a strong decline of B concentrations occur in ganders. In layers, T is further increased, E2 and P4 shows high levels, and, at the same time DHEA and T3 remain unchanged, while B and T4 decrease.

Achieving luster: prenuptial molt pattern predicts iridescent structural coloration in Blue-black Grassquits

Colors in feathers are produced by pigment deposition or by nanostructures within barbs or barbules. In the absence of pigments or nanostructures, light is scattered incoherently, producing white coloration. Honest advertisement models predict that ornamental colors evolve if they reliably signal individual properties such as viability, health, or nutritional state. In this study, we tested if (1) iridescent structural and (2) unpigmented plumage coloration signal male quality in the Blue-black Grassquit (Volatinia jacarina). During three reproductive seasons, we captured males and measured morphological variables and nuptial plumage coverage, and collected feathers for spectrometry. We found that saturation of the iridescent coloration was positively related to relative molting speed, indicating that males investing more in prenuptial molt also produced more saturated, UV-shifted plumage. Body condition was not related to brightness or saturation of the iridescent plumage, and no male morphological attributes were associated with the white underwing patch coloration or size. Our results suggest that patterns of molt, and possibly feather growth, may affect the organization of optical nanostructures responsible for iridescent coloration, and that the ontogeny of iridescent plumage ornaments must be taken into account in hypotheses concerning honest advertisement of such signals. Thus, coloration in this species may reliably reflect energy allocation to molting, constituting an honest indicator of male quality, life history decisions, or endocrine state. To our knowledge, this is the first report of natural variation in molting pattern being associated with sexually-selected structural color plumage.

Plumage colour acquisition and behaviour are associated with androgens in a phenotypically plastic tropical bird

Androgens regulate male reproductive behaviour and may be a mechanistic link between sexual signals and physiological condition. However, the role of these hormones in regulating prenuptial moult and male plumage signals of passerine birds is unclear. In the red-backed fairy-wren, Malurus melanocephalus, plumage colour is a sexually selected trait and males express three reproductive phenotypes: males can breed in bright red and black plumage or in dull brown plumage, or assume dull plumage and act as nonbreeding auxiliaries; each phenotype differs in parental and reproductive behaviour. We found that plasma androgen concentrations differed significantly between male phenotypes, with red/black breeding males having the highest levels and auxiliaries having the lowest levels across all nesting stages. These hormonal differences were also present during the prebreeding moult when nuptial plumage is acquired. Males also differed significantly in body condition during moult based on the phenotype acquired in the subsequent breeding season, with red/black breeding males being in the best condition and auxiliaries being in the poorest condition. Moreover, androgen concentrations were positively correlated with body condition during prenuptial moult. Thus, in this species, androgens may determine plumage coloration and provide a link between the expression of sexual signals and body condition.

Can long‐distance migratory birds adjust to the advancement of spring by shortening migration distance? The response of the pied flycatcher to latitudinal photoperiodic variation

AbstractMany organisms use day length as a cue for synchronizing their life cycles with seasonal changes in environmental productivity. Under rapid climate change, however, responses to day length may become maladaptive, and photo‐responsive organisms may only be able to evade increasingly unsuitable habitats if they can accommodate to a wide range of photoperiodic conditions. A previous experiment showed that the pied flycatcher,Ficedula hypoleuca, a Palaearctic‐Afrotropical migratory bird, would strongly advance the timing of spring migration and reproductive maturation if it shifted its wintering area from sub‐Saharan Africa to the Mediterranean region. However, it is unknown whether this marked response to latitudinal variation in photoperiodic conditions is continuous over the entire range of potential wintering areas, and if a shortening of migration distance would be an effective mechanism to adjust the timing of migration to rapidly changing climatic conditions. Here, we experimentally show that a moderate northward displacement of the pied flycatcher's current wintering grounds by 10° would result in a clear advancement of the termination of prenuptial moult and the initiation of spring migratory activity and gonadal growth. However, we found no further advancement under conditions simulating higher wintering latitudes, suggesting the existence of a critical photoperiodic threshold or a steep gradual response within a narrow geographical range between 10° and 20° northern latitude. Because habitat conditions in this area are deteriorating rapidly, the potential for pied flycatchers to adjust their life cycle to changing climatic conditions by shortening the migration distance may be limited in the future.