Age‐related reproductive success in dichromatic male Black Redstarts Phoenicurus ochruros: Why are yearlings handicapped?

Reproductive constraints on sexually mature, young birds may be the underlying cause for the evolution and maintenance of delayed plumage maturation in passerine birds. I examined whether yearling male Black Redstarts in non-definitive plumage are reproductively handicapped and explored possible reasons for it. Yearlings raised only half as many young per season as adults. Differential reproductive effort was excluded as a possible source for age-related reproductive success because yearlings and adults did not differ in the frequency of non-breeding or in paternal effort. The lower annual reproductive success of yearlings was attributed to their lower probability of obtaining (multiple) females and adult females. Males mated to adult females raised almost twice as many offspring as males mated to yearling females. I conclude that the proposed preconditions for the evolution and maintenance of delayed plumage maturation do still prevail in the Black Redstart.     

Populationsanteile und H?henverbreitung einj?hriger M?nnchen beim Hausrotschwanz (Phoenicurus ochruros) im Waldviertel, Nieder?sterreich

Zuammenfassung Im niederösterreichischen Waldviertel (200–700 m üNN) stieg der relative Anteil einjähriger von 17,9 auf 50% aller revierverteidigenden Hausrotschwanz- mit zunehmender Meereshöhe. Die Zunahme betrifft vor allem einjährige dercairei-Morphe, während dieparadoxus-Morphe in geringerer Anzahl und gleichförmiger als in allen Höhenbereichen auftrat.

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The portion of first-year males in breeding populations of Black Redstarts (Phoenicurus ochruros) and their altitudinal distribution in the Waldviertel, Lower Austria

Summary In 1986, between April 13 and July 5, breeding territories of male Black Redstarts were mapped in Lower Austria (200–700 m NN), with regard to their age (first-year or older) and the plumage types of first-year (femalelike or adult malelike). Due to a greater percentage of femalelike subadult (cairei-type) in higher altitudes, the proportion of first-year was found to increase from 17,9 per cent at 200–300 m to 50 per cent above 600 m. On the contrary adultlike first-year (paradoxus-type) are evenly distributed in smaller numbers over the whole altitudinal range.

     

Timing of events on the breeding grounds for five species of sympatric warblers

ABSTRACT On the breeding grounds, migratory birds have limited time to breed and molt before autumn migration. However, few studies of long‐distance migrants have examined the phenology of these events to determine what life‐history trade‐offs might result if these activities overlap. From 2000 to 2007, I used banding data to determine the timing of migration, breeding, and primary molt for Yellow Warblers (Dendroica petechia), Yellow‐rumped Warblers (D. coronata coronata), American Redstarts (Setophaga ruticilla), Ovenbirds (Seiurus aurocapilla), and Canada Warblers (Wilsonia canadensis) at a study site in Alberta, Canada. Hatching date did not differ among species (P= 0.63), with means ranging from 27 June to 3 July. All species began primary molt between 12 July and 18 July, near the expected fledging date of offspring, and therefore all species exhibited overlap between postfledging parental care and molt. The duration of primary molt ranged from 28 d for Canada Warblers to 69 d for Yellow‐rumped Warblers. Yellow Warblers, Yellow‐rumped Warblers, and American Redstarts began autumn migration having completed about 50% of their primary molt. However, Ovenbirds departed when 21% of molt was complete, and Canada Warblers departed 2 d after completing molt. For all five species of warblers, molt did not overlap with nest‐bound breeding activities. However, molt did overlap with both postfledging care and migration. This suggests that initiating migration as soon as possible is important, possibly because earlier arrival on the wintering grounds may improve access to high quality winter habitat. Overall, warblers may maximize individual fitness by combining life‐history events that result in overlapping portions of the breeding cycle, molt, and migration.     

Complex postbreeding molt strategies in a songbird migrating along the East Asian Flyway,the Pallas’s Grasshopper Warbler Locustella certhiola

Molt strategies have received relatively little attention in current ornithology, and knowledge concerning the evolution, variability and extent of molt is sparse in many bird species. This is especially true for East Asian Locustella species where assumptions on molt patterns are based on incomplete information. We provide evidence indicating a complex postbreeding molt strategy and variable molt extent among the Pallas's Grasshopper Warbler Locustella certhiola, based on data from six ringing sites situated along its flyway from the breeding grounds to the wintering areas. Detailed study revealed for the first time that in most individuals wing feather molt proceeds from the center both toward the body and the wing‐tip, a molt pattern known as divergent molt (which is rare among Palearctic passerines). In the Russian Far East, where both breeding birds and passage migrants occur, a third of the adult birds were molting in late summer. In Central Siberia, at the northwestern limit of its distribution, adult individuals commenced their primary molt partly divergently and partly with unknown sequence. During migration in Mongolia, only descendantly (i.e., from the body toward the wing‐tip) molting birds were observed, while further south in Korea, Hong Kong, and Thailand the proportion of potential eccentric and divergent feather renewal was not identifiable since the renewed feathers were already fully grown as expected. We found an increase in the mean number of molted primaries during the progress of the autumn migration. Moderate body mass levels and low‐fat and muscle scores were observed in molting adult birds, without any remarkable increase in the later season. According to optimality models, we suggest that an extremely short season of high food abundance in tall grass habitats and a largely overland route allow autumn migration with low fuel loads combined with molt migration in at least a part of the population. This study highlights the importance of further studying molt strategy as well as stopover behavior decisions and the trade‐offs among migratory birds that are now facing a panoply of anthropogenic threats along their flyways.     

The Costs of Male Display and Delayed Plumage Maturation in the Satin Bowerbird (Ptilonorhynchus violaceus)

The costs associated with the evolution of male display traits has attracted much attention in regard to the type of traits that evolve and the timing of their expression. We investigate these costs by using testosterone implants to alter the development of male display traits in the satin bowerbird (Ptilonorhynchus violaceus). Testosterone implants advanced the development of adult display traits in males with juvenile plumage. We then measured the costs of early expression of these characters in treated males. Testosterone implants caused young males to (1) become more involved in behavior normally carried out by older males when visiting bowers, (2) build bowers, and (3) molt prematurely into an adult plumage. No difference was observed between the testosterone-treated birds and controls in return rate or condition the year following treatment, suggesting that the physical costs of display are not high. We found that males in adult plumage are tolerated less, and are not displayed to as frequently as juvenile plumage males at the bowers of established bower holders. The causes of delayed plumage maturation are most consistent with the facilitated-learning hypothesis. That is, early development of adult characteristics reduces the opportunity of young males to learn display behavior critical for later reproductive success.     

Demographic variation in timing and intensity of feather molt in migratory Fork‐tailed Flycatchers (Tyrannus s. savana)

Understanding the annual cycle of migratory birds is imperative for evaluating the evolution of life‐history strategies and developing effective conservation strategies. Yet, we still know little about the annual cycle of migratory birds that breed at south‐temperate latitudes of South America. We aged, sexed, and determined the progression and intensity of body, remige, and rectrix molt of migratory Fork‐tailed Flycatchers (Tyrannus s. savana) at breeding sites in southern South America and at wintering sites in northern South America. Molt of both body and flight feathers occurred primarily during the winter. In early winter, a similar proportion of young and adult flycatchers molted remiges and rectrices, but remige molt intensity (number of remiges molting) was greater and primary molt progression (mean primary feather molting) more advanced in adults. In late winter, remige molt intensity and primary molt progression did not differ between age groups. We found no difference between males and females either in the proportion of individuals molting in winter or in the intensity or progress of remige molt. Our results suggest that the nominate subspecies of Fork‐tailed Flycatcher undergoes one complete, annual molt on the wintering grounds, and represents the first comprehensive evaluation of molt timing of a migratory New World flycatcher that overwinters in the tropics. Given that breeding, molt, and migration represent three key events in the annual cycle of migratory birds, knowledge of the timing of these events is the first step toward understanding the possible tradeoffs migratory birds face throughout the year.     

Invertebrate availability and vegetation characteristics explain use of nonnesting cover types by mature‐forest songbirds during the postfledging period

ABSTRACT Some species of mature‐forest‐nesting songbirds use regenerating clearcuts and forested wetlands during the postfledging period (between nesting and migration). Relatively dense vegetation structure and abundant food resources in non‐mature‐forest cover types have been hypothesized to explain this phenomenon. We examined the relative importance of vegetation structure and invertebrate availability on use of nonnesting cover types by adult and hatch‐year Ovenbirds (Seiurus aurocapilla) and American Redstarts (Setophaga ruticilla) during the postfledging period of 2009 in northern Minnesota. We used mist nets to sample bird use of forested wetlands and regenerating clearcuts of three age groups: 1–6, 7–12, and 16–19 yr after harvest. We modeled captures of birds using vegetation characteristics and invertebrate availability sampled around nets as explanatory variables. For all birds studied, captures were best explained by food availability and secondarily by vegetation characteristics including litter depth and woody debris for Ovenbirds and canopy height for American Redstarts. Shrub‐level invertebrate availability received a cumulative weight of 0.74–0.99 in Akaike's information criterion corrected ranked models for adult and hatch‐year birds of both species. Vegetation density and variation in vegetation density explained almost no variation in captures of either species. We conclude that both invertebrate availability and some vegetation characteristics influence use of nonnesting cover types by Ovenbirds and American Redstarts during the postfledging period, but that invertebrate availability is generally the stronger predictor of that use.     

Sources of distinctness of juvenile plumage in Western Palearctic passerines

Juveniles of many avian species possess a spotted or mottled body plumage that is visually distinct from the plumage of adults. In other species, however, juveniles fledge with a body plumage that is just a pale representation of adult female plumage. The reasons for this variation are poorly understood. Several hypotheses concerning social (parent–offspring, adult–juvenile, juvenile–juvenile), ecological (predation risk) and physiological (costs of plumage development) implications of juvenile body plumage are presented in relation to predictions concerning associations with certain ecological and life‐history attributes of avian species. In the present study, we conduct a phylogenetically corrected comparative analysis of Western Palearctic passerines looking for sources of variation in the incidence of distinct and adult‐like juvenile body plumages. We scored plumages based on plates in the Handbook of the Birds of the Western Palearctic (Cramp & Perrins, 1988–1994; Oxford University Press) (HBWP) and entered body mass, migratory habits, habitat, nestling diet, breeding dispersion, gregariousness, duration of the nestling period, type of nest, conspicuousness of female plumage, and sexual dimorphism as explanatory variables, as presented in HBWP, in phylogenetic generalized least square regression analyses. One‐third of the species presented distinct juvenile body plumages, which lasted on average for the first 2 months of life. Body mass, conspicuousness of female plumage, migratory habits, and habitat were significantly associated with interspecific variation in distinctness of juvenile plumage, with smaller species, more conspicuous species, migrants, and species from forested habitats showing distinct juvenile plumages with higher frequency. The phylogenetic signal was moderately high. Assuming that conspicuous adult plumage is costlier to produce than distinct juvenile body plumage (pigments, conspicuousness), the need to acquire social status among juveniles before the winter may explain the more adult‐like plumage in resident species because juveniles will probably compete with individuals that they may have known during their first months of life. On the other hand, migrant juveniles may compete with a different set of individuals in winter quarters and can use savings in resources necessary for developing adult‐like plumages to improve migration capacity by allocating resources to other functions. The association with habitat could be related to juveniles in open habitats participating in more extended interactions with other juveniles than in forested habitats where lower visibility may reduce the capacity to detect or respond to signals from juvenile conspecifics. More studies on this possibly crucial life stage are needed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 440–454.     

Plumage coloration and ornamentation as predictors of nest survival and number of young fledged in Yellow Warblers

ABSTRACT Previous studies of Yellow Warblers (Setophaga petechia) indicate that males with heavy melanin‐based brown breast streaking are more territorial and obtain more extra‐pair copulations than lightly streaked males that provide more parental care and are cuckolded more often. Because carotenoid‐based plumage characters can also signal quality in male songbirds, we examined relationships between yellow carotenoid‐based breast coloration and brown breast streaking and measures of the reproductive performance of male and female Yellow Warblers in northwestern Ohio from April to June 2007–2008. We used an information theoretic approach to examine relationships between reproductive measures (daily nest survival rate and number of young fledged) and brown breast streaking, yellow breast coloration, extent of prealternate wing molt, age, year, and first‐egg date. Males with paler yellow breast plumage (Beta hat =–0.213) and those with more brown breast streaking (Beta hat =–0.000036) fledged fewer young, but we found no associations between number of young fledged per male and age, year, or extent of prealternate wing molt. In addition, none of the measured variables was associated with the number of young fledged per female. Age was positively associated with daily nest survival rates for male Yellow Warblers (Beta hat = 1.127), but we found no association between daily nest survival rate and brown breast streaking, yellow breast coloration, extent of prealternate wing molt, age, year, and first‐egg date. None of the measured variables was associated with daily nest survival for females (null model was top‐ranked). Because brown breast streaking and the intensity of yellow breast coloration of male Yellow Warblers were negatively related to number of young fledged, we suggest that these plumage characteristics function differently in male–female and male–male interactions; yellow breast coloration signals age and reproductive experience, whereas brown breast streaking signals degree of territoriality or parental care.     

Age‐dependent relationships between coloration and reproduction in a species exhibiting delayed plumage maturation in females

Males of many bird species exhibit delayed plumage maturation (DPM), a condition in which young individuals display an immature plumage. Several adaptive hypotheses have been suggested for the signaling utility of DPM in males. Tree swallows Tachycineta bicolor, however, are one of the few bird species to exhibit DPM in females, but not in males. Few studies have focused on the age‐dependent signaling function of female plumage traits due to the uncommon nature of DPM in females. Therefore, we used reflectance spectrometry and scanning electron microscopy of sub‐adult (melanin‐based brown) and adult (iridescent‐blue structural) female tree swallows to characterize plumage coloration. Next, we asked whether variation in plumage coloration in females reflects condition and reproductive performance between and within age classes. We found that older females were in better body condition and laid eggs earlier in the season compared to young females; however, average egg mass and reproductive success (number of offspring fledged and offspring condition) did not differ between age classes. There were significant relationships indicating that young females with more‐ornamented (darker brown melanin) plumage laid smaller eggs, but hatched eggs earlier in the season leading to nestlings in better condition compared to less‐ornamented young females. Older females that were more ornamented (brighter, greater blue chroma, and lower hues in iridescent plumage) laid heavier eggs, but ornamentation was negatively associated with immune function, health, and reproductive success. Together, these data suggest that female ornamentation reflects reproductive performance and that there are complicated relationships between plumage coloration, condition, and reproductive performance that ultimately influence reproductive success.     

Conditions on the Mexican moulting grounds influence feather colour and carotenoids in Bullock's orioles (Icterus bullockii)

Carotenoid‐based plumage coloration plays a critical role for both inter‐ and intrasexual communication. Habitat and diet during molt can have important consequences for the development of the ornamental signals used in these contexts. When molt occurs away from the breeding grounds (e.g., pre‐alternate molt on the wintering grounds, or stopover molt), discerning the influence of habitat and diet can be particularly important, as these effects may result in important carryover effects that influence territory acquisition or mate choice in subsequent seasons. Several species of songbirds in western North America, including the Bullock's oriole (Icterus bullockii), migrate from the breeding grounds to undergo a complete prebasic (post‐breeding) molt at a stopover site in the region affected by the Mexican monsoon climate pattern. This strategy appears to have evolved several times independently in response to the harsh, food‐limited late‐summer conditions in the arid West, which contrast strongly with the high productivity driven by heavy rains that is characteristic of the Mexican monsoon region. Within this region, individuals may be able to optimize plumage coloration by molting in favourable areas characterized by high resource abundance. We used stable isotope analysis (δ¹³C, δ¹⁵N) to ask whether the diet and molt habitat/location of Bullock's orioles influenced their expression of carotenoid‐based plumage coloration as well as plumage carotenoid content and composition. Bullock's orioles with lower feather δ¹⁵N values acquired more colorful plumage (orange‐shifted hue) but had feathers with lower total carotenoid concentration, lower zeaxanthin concentration, and marginally lower canthaxanthin and lutein concentration. Examining factors occurring throughout the annual cycle are critical for understanding evolutionary and ecological processes. Here, we demonstrate that conditions experienced during a stopover molt, occurring hundreds to thousands of kilometers from the breeding grounds, influence the production of ornamental plumage coloration, which may carryover to influence inter‐ and intrasexual signaling in subsequent seasons.     

Vergleichende Untersuchung von Jugendentwicklung und Zugverhalten bei Garten- und Hausrotschwanz,Phoenicurus phoenicurus undPh. ochruros

Zusammenfassung An je 12 handaufgezogenen Garten- und Hausrotschwänzen wurden Jugendmauser, Fettdeposition und Zugunruhe erfaßt und mit entsprechenden Daten freilebender Artgenossen verglichen. Die Daten stimmen gut mit denen freilebender Artgenossen überein. Beim Gartenrotschwanz setzt die Jugendmauser in frühem Alter ein und verläuft rasch, und Fettdeposition und Zugunruhe beginnen, ehe die Mauser beendet wird. Er zeigt das typische Bild eines frühwegziehenden Interkontinentalziehers. Beim Hausrotschwanz hingegen wird die Jugendmauser trotz ihrer großen Länge beendet, bevor Fettdeposition und Zugunruhe einsetzen, und beide haben geringeren Umfang. Er bietet das Bild eines weniger ausgeprägten Ziehers und eines Intrakontinentalziehers. Beide Arten zeigen ihre großen Unterschiede in Jugendentwicklung und Zugverhalten in weitgehend entsprechenden Versuchsbedingungen, vor allem unter ganz ähnlichen photoperiodischen Bedingungen. Das spricht dafür, daß diese Unterschiede einer stark endogenen, wahrscheinlich einer stark genetischen Kontrolle unterliegen. Der Gartenrotschwanz — als Weitstrekkenzieher — entwickelt viel, der Hausrotschwanz — als Kurz- bis Mittelstreckenzieher — weit weniger Nachtunruhe. Die Zeit, in der Nachtunruhe produziert wird, stimmt bei beiden Arten sehr gut mit der Zugzeit überein. Die Nachtunruhe ist daher bei beiden Arten typische Zugunruhe. Die verschiedenen gemessenen Zugunruhewerte stimmen z. T. sehr gut mit den zurückzulegenden Zugstrecken überein, z. T. zeigen sie erhebliche Abweichungen. Prinzipiell lassen sich die Daten jedoch alle gut im Sinne endogener Zug-Zeitprogramme interpretieren, wie sie früher für Sylviiden nachgewiesen wurden. Demnach lassen sich die Vorstellungen endogener Zug-Zeitprogramme nunmehr auf eine weitere Vogelgruppe, nämlich die kleinen Drosselvögel, ausdehnen.

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A comparative study of juvenile development, migratory restlessness and migratory behaviour in the Redstart and Black Redstart

Summary Juvenile development, migratory disposition and migratory activity were investigated in Redstarts and Black Redstarts. In each 12 hand-raised individuals, patterns of juvenile moult, fat deposition and migratory restlessness were determined and compared with corresponding data from freeliving conspecifics. There was good agreement between the data from hand-raised and feral birds. In the Redstart, juvenile moult starts early and proceeds rapidly and fat deposition and migratory restlessness start before the moult has ended. The Redstart is typical for an early departing intercontinental migrant. In the Black Redstart, however, the moult, despite its long duration was clearly separated from the migratory events, and fat deposition and migratory restlessness are both less pronounced than in the Redstart. The patterns of the Black Redstart were typical for a less pronounced or an intracontinental migrant. Both species produced the large differences in juvenile development and migratory behaviour in widely corresponding experimental conditions, above all in very similar photoperiodic conditions. This suggests that these differences were based to a high degree on endogenous, probably genetic factors. The Redstart, as a long-distance migrant, developed a lot, and the Black Redstart, a short- to middle-distance migrant, far less nocturnal restlessness. In both species the period in which nocturnal restlessness developed corresponded very well with the migratory season. Thus this restlessness was typical migratory restlessness. The various measurements of restlessness obtained corresponded in part with the distances covered during migration although there were substantial deviations. In principle, however, all the data can well be interpreted in terms of endogenous time programs for migration as they have been formerly demonstrated for Sylviid species. Thus the concept of these programs can now be extended to another bird group — to small Turdiid species.

     

Post-fledging interactions between the Common Cuckoo Cuculus canorus and its cavity-nesting Common Redstart Phoenicurus phoenicurus host

Brood parasite–host interactions during the incubation and nestling stages have been well studied, but the post-fledging period remains virtually unknown. Using radiotracking, we provide the first detailed data on post-fledging interactions between the Common Cuckoo Cuculus canorus and its only regular cavity-nesting host, the Common Redstart Phoenicurus phoenicurus. Cuckoos raised alone (‘solitary’) fledged at higher mass, with higher wing and tarsus length and started to fly at a younger age than Cuckoos raised alongside young Redstarts (‘mixed’). However, a further 23 fledging and post-fledging parameters measured at five pre-determined times (fledging, first-flight, predation, starvation, independence) did not differ between solitary and mixed Cuckoos. In addition, none of the parameters measured during the post-fledging period (growth, dispersal distances, number of flights) differed between solitary and mixed Cuckoos. Redstart fledglings from non-parasitized broods (‘solitary’) showed generally similar fledging and post-fledging parameters to fledglings reared alongside a Cuckoo (‘mixed’). Surprisingly, there were no significant differences in post-fledging predation rate, starvation or overall survival rates between mixed and solitary Cuckoos or mixed and solitary Redstarts. Thus, during the post-fledging period, mixed Cuckoo fledglings successfully compensated for the poorer performance experienced during the nestling stage whereas mixed and solitary Redstarts did not differ in any measured parameters. This suggests that the regular occurrence of mixed broods in this host–parasite system – which is unique among the many Cuckoo hosts – is evolutionarily stable for both hosts and parasites.     

Melanin‐based coloration covaries with fluctuating asymmetry,nutritional state and physiological stress response in common snipe

Adaptative significance of melanin‐based coloration in birds remains poorly recognized. It has been suggested that genes responsible for melanin synthesis may have pleiotropic effect on several physiological and behavioural functions, including immune defence. For this reason, we could expect that the expression of melanin‐based plumage coloration should covary with different condition‐related phenotypic traits via regulation of pathogen/parasite resistance. The aim of this study was to test this hypothesis in common snipe Gallinago gallinago, a species that exhibits conspicuous variation in the black eumelanism of underwing plumage. The study was conducted in central Poland, where common snipe were captured during autumn migration. We found that after accounting for the effects of age, sex and date of capture, underwing coloration correlated with nutritional state of snipes, as more extensively melanised individuals had higher plasma concentrations of triglycerides, total protein and albumin. Dark underwing coloration was also associated with lower heterophil/lymphocyte (H/L) ratio, suggesting better resistance of eumelanic individuals to physiological stress. Finally, adult males with darker underwings had lower asymmetry in wing shape (wingtip convexity), which indicated their higher developmental stability. In conclusion, melanin‐based coloration may be considered an honest indicator of phenotypic quality in common snipe.     

Effects of migration distance on life history strategies of Western and Semipalmated sandpipers in Perú

Migration distances of shorebird species correlate with life history strategies. To assess age‐specific migratory preparation and adult wing‐molt strategies, we studied Western Sandpipers (Calidris mauri) and Semipalmated Sandpipers (C. pusilla) with different migration routes at the Paracas National Reserve in Perú, one of the most austral non‐breeding areas for these sandpipers, from 2012 to 2015. Western Sandpipers breed near the Bering Sea, ~11,000 km from Paracas. Semipalmated Sandpiper populations at Paracas are a mixture of short‐billed birds from western Arctic breeding sites, plus long‐billed birds from eastern sites, ~8000 km distant. Adults of both species arrive in October with primary feathers already partially renewed so wing molt starts at sites further north. Semipalmated Sandpipers with longer bills completed wing molt later than shorter billed birds. Adults of both species prepared for migration in February and March. No juvenile Western Sandpipers prepared for migration, confirming the “slow” over‐summering life history strategy of more southerly non‐breeding populations. Juvenile Semipalmated Sandpipers showed bimodality in strategies. Most showed no migratory preparation, but, during three non‐breeding periods, from 27% to 31% fattened, molted, and partially replaced outer primaries during the pre‐migratory period. Juveniles with longer culmens were heavier and tended to have more alternate plumage. Juveniles that were partially molting primaries had longer culmens and more alternate plumage. Juvenile Semipalmated Sandpipers from eastern‐breeding populations thus have a higher propensity for a fast life history strategy, and western birds a slow one, at this non‐breeding site in Peru. Western‐breeding Semipalmated Sandpiper populations thus resemble Western Sandpipers, suggesting a common, possibly distance‐related, effect on life history strategy.     

Sexually selected male plumage color is testosterone dependent in a tropical passerine bird, the red-backed fairy-wren (Malurus melanocephalus)

Background

Sexual 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 Findings

We 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/Significance

We 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.     

Temporal and spatial patterns of flight and body feather molt of Bank,Barn, and Cliff swallows in North and South America

Molt is energetically demanding and various molt strategies (i.e., molt series, duration, intensity, timing, and location) have evolved to reduce the negative fitness consequences of this process. As such, molt varies considerably among species. Identifying where and when specific feathers are molted is also crucial to inform species‐specific studies using stable isotope markers to assign individuals to geographical regions where they molt. Using museum specimens, we examined the molt of three species of migratory swallows in the Americas: Bank Swallows (Riparia riparia), Barn Swallows (Hirundo rustica), and Cliff Swallows (Petrochelidon pyrrhonota). All three species have one primary and two secondary molt series. Bank and Cliff swallows had one rectrix molt series, and Barn Swallows molted the outer rectrix (R6) separately from the inner five rectrices (R1‐5). All three species have a relatively long flight feather molt duration (i.e., 140–183 days) and low molt intensity. Barn Swallows initiated flight feather molt in the fall, about 2 months later than Bank and Cliff swallows. Barn Swallows likely delay molt because of constraints associated with double brooding. For all three species, molt started with the primaries and inner secondaries and was closely followed by the rectrices and, finally, the outer secondaries. For those that began and then interrupted molt either in breeding areas or during fall migration, the first feathers molted were predominantly S8 and P1. All three species underwent body molt throughout the year, but most individuals molted their body plumage in wintering areas. We recommend that the most appropriate feathers for stable isotope research examining migratory connectivity and habitat use are either R2‐R4 or S2‐S4.     

GPS‐tracking reveals non‐breeding locations and apparent molt migration of a Black‐headed Grosbeak

Black‐headed Grosbeaks (Pheucticus melanocephalus) have been observed to undergo prebasic molt during fall in the North American Monsoon region of the southwestern United States and northwestern Mexico, but it is unknown whether molt migration is pervasive across populations of the species. During the 2014 breeding season, we GPS‐tagged (where GPS is global positioning system) nine adult Black‐headed Grosbeaks in Yosemite National Park with archival GPS tags to determine specific locations where grosbeaks breeding in Yosemite spent portions of the non‐breeding season, and to assess whether those locations were consistent with molt migration. On 2 June 2015, one of these birds, a male GPS‐tagged on 19 June 2014, was recaptured with its GPS unit still attached. Data downloaded from the unit revealed that, by 20 August 2014, the bird had moved 1300 km from Yosemite National Park to Sonora, Mexico, where it remained until at least 15 October 2014. By 24 November 2014, the grosbeak had moved >1300 m from Sonora to the Michoacán‐Jalisco border region, where it remained until the last GPS‐determined location was obtained on 24 March 2015. The seasonal timing of these movements and the length of stay in Sonora are consistent with the expected behavior of a molt‐migrating bird. Remote‐sensed enhanced vegetation index (EVI) data indicated that the grosbeak arrived in the monsoon region near the area's annual peak in EVI, and then, as the index was declining sharply, departed for the Michoacán‐Jalisco region, where the index also declined during the same period, but substantially less so than in Sonora. Climate change in the coming decades is expected to delay the annual onset of the monsoon while also accelerating the initiation of arid, summer‐like conditions throughout much of western North America, possibly yielding a temporal mismatch between fall migration and the monsoon‐driven conditions that may be critical for molt‐migrating birds.     

Males in seemingly female‐like plumage do not mimic females: UV reflectance reveals temporal cryptic dimorphism in a manakin species exhibiting delayed plumage maturation

Manakins (Pipridae) are neotropical birds that usually exhibit delayed plumage maturation (DPM). Thus, while plumage of most adult male manakins is brightly conspicuous, subadult males and females are basically dull‐olive green. Although sexual dichromatism in some bird species may be evident only through UV reflectance, this phenomenon, known as hidden sexual dichromatism, has not been previously studied in manakins to compare subadult males and females. Within this framework, we carried out spectrophotometric analyses in searching for hidden sexual dichromatism in the white‐bearded manakin Manacus manacus, through comparison of UV spectra in females and subadult males in green plumage. Our results revealed UV reflectance in both sexes in green plumage. Moreover, we found UV spectral differences in homologous color patches between sexes, particularly at belly. Since the observed differences may allow intraspecific sex recognition of individuals in green plumage, our results do not support the female‐mimicry hypothesis to explain delayed plumage maturation in the white‐bearded manakin. Although our findings dismiss the female mimicry hypothesis, we cannot state whether these results support the non‐mutually exclusive cryptic and status signaling hypotheses. We propose then, that dull coloration of subadult males may serve both as a cryptic trait and to limit the energetic costs of acquiring the adult plumage before sexual maturity. Meanwhile, differential UV color traits between sexes in green plumage may allow adult males to avoid unnecessary energy expenditures in courtship displays in the presence of males near leks, and to selectively focus their the courtship displays on females. In accordance with the status signaling hypothesis, subadult males can be recognized both as males and subordinates and consequently may practice courtship displays without suffering aggressions by adult males. Our results highlight the importance to include a wider range of spectrophotometric information analyses for testing hypotheses regarding delayed plumage maturation.     

Age structure and age‐related differences in molt status and fuel deposition of Dunlins during the nonbreeding season at Chongming Dongtan in east China

ABSTRACT Although most shorebirds exhibit deferred migration and deferred breeding during their first summer, Dunlins (Calidris alpina) migrate to breeding areas and breed during their first summer. First‐year and adult Dunlins should, therefore, have similar fueling and molt patterns if energetic and physiological constraints are responsible for deferred migration. From 2006 to 2008, we examined the age structure of Dunlins during the nonbreeding season at Chongming Dongtan, an estuarine wetland in the Yangtze River estuary in east China, and examined the effects of date, age, and molt status on fuel deposition during migration and during the winter. The Dunlin population at Chongming Dongtan was composed primarily of first‐year birds. Most adults and first‐year birds arrived together in late August. Regression analyses indicated that age, date, and molt status affected fuel deposition (as indicated by body mass) of Dunlins. Adults had significantly greater fuel deposits than first‐year Dunlins near the end of northward migration (May: adults 70.8 ± 6.4 g, first‐year 63.8 ± 8.0 g) and at the start of southward migration (September: adults 50.2 ± 6.1 g, first‐year 47.2 ± 4.9 g). Adults also had significantly higher fuel deposition rates than first‐year Dunlins during northward migration. Nonetheless, first‐year Dunlins migrate and breed in their first summer. Thus, other factors, such as migration distance and body size, may be more important in determining if first‐year shorebirds defer migration during their first spring and summer. During boreal spring and autumn, first‐year Dunlins in active body molt had greater body mass than those that had not initiated body molt or those in suspended molt, and premigratory fuel deposits for northward migration were greatest after prealternate molt was completed. These results suggest that body molt requires additional fuel deposits and imposes a constraint on fuel deposition for migratory flights.