Prebasic Molt Research Articles

Molts and Plumages of Ducks (Anatinae): An Evaluation of Pyle (2005)

Ducks are unusual because males of most Holarctic species acquire brightly-pigmented breeding plumages in autumn and winter and not spring as with most other birds. Based on an evaluation of molt phenology of ducks under the system of molt terminology invented by Humphrey and Parkes (1959), Pyle (2005) concluded that, for most North American duck species, the molt that produces bright plumages is basic in nature, the molt that typically produces a short-lived, dull “eclipse” plumage in spring or summer is alternate in nature, and the annual wing molt that largely occurs between these two molts is part of the subsequent recharacterized prebasic molt. For various reasons, it appears more likely that the timing of conventional prealternate molts in most Holarctic duck species has accelerated forward over time to autumn and winter in response to selection for early courtship and pairing, with resulting changes in the duration of alternate and basic plumages. These reasons include acceleration in the timing of the prealternate molt in sea ducks after they reach the age of first breeding. An accelerated timing of prealternate molts can explain why certain ducks exhibit an ephemeral, limited first prebasic molt, which appears to be a lost molt and not a virtually nonexistent, unique preformative molt as maintained by Pyle (2005). The accelerated timing hypothesis also can explain why the extent of prealternate molts may have become more complete and the extent of prebasic molts may have become less complete in certain ducks. The accelerated timing hypothesis appears more parsimonious from an evolutionary perspective than the system of Pyle (2005), and does not require the categorical rejection of plumage color and pattern in molt homology analyses, as advocated by Pyle (2005).

Molt Patterns, Biometrics, and Age and Gender Classification of Landbirds on Saipan, Northern Mariana Islands

Abstract Molt strategies and plumage development by age and gender are poorly understood for resident tropical landbirds. We used data from six banding stations on Saipan, Northern Mariana Islands, and examination of 267 museum specimens to describe patterns of molt and to establish criteria for assessing age and gender of nine native resident landbird species on the island. Feather replacement sequences in the majority of Saipan's resident landbirds were typical of related species. Preformative molts occurred in at least eight of the nine species; these and definitive prebasic molts were incomplete to complete, and prealternate molt appeared to be absent for all species. We developed criteria for classifying gender of sexually dimorphic species using plumage cues, presence of brood patch or cloacal protuberance, and biometrics. We confirmed whether or not both brood patch and cloacal protuberance were reliable indicators of gender during breeding in monomorphic species. Distinct periods or seasons of mol...

Comparison of adrenocortical responses to acute stress in lowland and highland Eurasian tree sparrows (Passer montanus): similar patterns during the breeding, but different during the prebasic molt

Previous studies indicate most free-living avian species in both extreme and temperate environments seasonally modulate the adrenocortical responses to acute stress, and those breeding in harsh environments always express reduced adrenocortical responses, which may allow them to obtain maximal reproductive success. However, recent investigations showing a human commensal species, house sparrows (Passer domesticus), expressed similar corticosterone (CORT) responses in both benign and harsh environments. In this study, focusing on another human commensal species, Eurasian tree sparrows (P. montanus), we examined the adrenocortical response to acute stress in lowland populations, among the early and late breeding, the prebasic molt, and the wintering stages, and compared them with previously published data from populations on the Tibetan Plateau. Our results show: (1) similar to highland Eurasian tree sparrows, lowland populations show no differences in baseline CORT levels among life history stages, and the stress-induced CORT (maximal CORT, total and corrected integrated CORT) levels are lower during the early breeding and the prebasic molt stages than those in the late breeding and the wintering stages; (2) highland Eurasian tree sparrows show stronger adrenocortical responses during the prebasic molt stage than lowland populations, whereas there are no differences between the early and the breeding stages (except for maximal CORT). Our results suggest that Eurasian tree sparrows from both harsh and benign environments have similar patterns of adrenocortical responses in the breeding stage, whereas they are different in the prebasic molt stage. In highland birds, the increased maximal CORT levels during the late breeding and the small increases in adrenocortical responses during the prebasic molt are interesting but remain unexplained.

Testing Competing Hypotheses For Chronology and Intensity of Lesser Scaup Molt During Winter and Spring Migration

Abstract. We examined chronology and intensity of molt and their relationships to nutrient reserves (lipid and protein) of Lesser Scaup (Aythya affinis) to test predictions of two competing hypotheses. The “staggered cost” hypothesis states that contour-feather molt is nutritionally costly and should not occur during nutritionally costly periods of the annual cycle unless adequate nutrients are available. The “breeding plumage” hypothesis states that prealternate molt must be complete prior to nesting, regardless of nutrient availability. Males and females were completing prebasic molt during winter (Louisiana) and had similar molt intensities. Females underwent prealternate molt during spring migration (Illinois and Minnesota) and prebreeding (Manitoba) periods; 53% and 93% of females were in moderate to heavy molt in Minnesota and Manitoba, respectively, despite experiencing other substantial nutritional costs. Intensity of prealternate molt was not correlated with lipid reserves even though females, on...

Implications of Prebasic and a Previously Undescribed Prealternate Molt For Aging Rusty Blackbirds

Abstract. Aging birds often relies on differences in plumage between immatures and adults, and understanding these patterns can improve our ability to discern demographic patterns within populations. We investigated patterns of prebasic molt of the Rusty Blackbird (Euphagus carolinus) in fall at Whitehorse, Yukon Territory, and developed a new technique for aging based on characteristics of the head plumage acquired during prebasic molt. Furthermore, we investigated the possibility of a prealternate molt among wintering Rusty Blackbirds on the basis of captures in Mississippi and museum specimens from across the species' winter range. Finally, we examined how a prealternate molt might affect the aging of winter birds by plumage. Rusty Blackbirds completed their prebasic molt by the end of September, and immature birds had a more prominent eye ring and a paler chin than adults, allowing a reliable age determination. Previously, the Rusty Blackbird was thought to attain its breeding plumage through feather ...

Variation in deuterium (δD) signatures of Yellow Warbler Dendroica petechia feathers grown on breeding and wintering grounds

Stable isotope analysis of feathers may help track populations of migratory birds throughout their annual cycle. We use deuterium isotope ratios in feathers (δDf) to confirm that Yellow Warblers Dendroica petechia go through both a prebasic molt on or near their breeding grounds and a prealternate molt on their wintering grounds, and evaluate whether δD analysis of different feather types can help assign birds to breeding and winter origins. We show that primary coverts and tail feathers have δDf values that reflect long-term average deuterium isotope ratios in precipitation (δDP) on the breeding grounds. Most greater coverts and crown feathers, however, have δDf values that are more enriched than predicted based on the δDP of their wintering distribution. We also found significant interannual variation in the δDf values of tail feathers, greater coverts and crown feathers. Interannual variation in δDf of winter-grown feathers may be explained by El Nino/La Nina conditions that influence rainfall and thus δDP; feathers were more depleted following an El Nino year (2005) than a La Nina year (2006). Gender had no effect on the δDf values of any feathers sampled. Age influenced the δDf values of crown feathers but not greater coverts. This study therefore confirms that δDf of summer-grown feathers can be used to identify breeding locations, but suggests that the ability to use δDf of winter-grown feathers to assign individuals to winter locations is limited by weak spatial gradients and temporal variation in δDP.

Feather isotope analysis discriminates age-classes of Western, Least, and Semipalmated sandpipers when plumage methods are unreliable

Avian age-class discrimination is typically based on the completeness of the first prebasic molt. In several calidrid sandpiper species, juvenal flight feathers grown on Arctic breeding grounds are retained through the first three migrations. Thereafter, flight feathers are grown annually at temperate migratory stopover sites during the fall or on the subtropical wintering grounds. Standard methods for distinguishing age classes of sandpipers rely on a combination of traits, including body plumage, coloration of protected inner median covert edges, and extent of flight feather wear. We tested the ability of stable hydrogen isotope ratios in flight feathers (Df )t o distinguish young birds in their first winter through second fall from older adults in three calidrid sandpiper species, Western (Calidris mauri), Least (C. minutilla), and Semipalmated (C. pusilla) sandpipers. We compared the apparent reliability of the isotope approach to that of plumage-based aging. The large expected differences in Df values of flight feathers grown at Arctic versus non-Arctic latitudes enabled use of this technique to discriminate between age-classes. We determined Df values of known Arctic-grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather Df values of southward-migrating adults showed bimodal distributions for all three species. Negative values overlapped with species-specific juvenile values, identifying putative second fall birds with high-latitude grown juvenal feathers retained from the previous year. The more positive values identified older adults who grew their feathers at mid- and low latitudes. Importantly, Df analysis successfully identified first-winter and second-fall birds not detected by plumage-based aging. Flight feather wear alone was a poor basis for age classification because scores overlapped extensively between putative second fall birds and older adults. Flight feather hydrogen isotope analysis enables more definitive assignment of age classes when standard plumage methods are unreliable. RESUMEN. El ande is´

PREBASIC MOLT OF BLACK-CAPPED AND WHITE-EYED VIREOS: EFFECTS OF BREEDING SITE AND THE EL NIÑO–SOUTHERN OSCILLATION

Abstract Ecological factors on the breeding grounds are expected to have a relatively large effect on the timing of molt in migratory birds breeding at middle latitudes (e.g., southern North America), because constraints on the time available to molt are probably weak. We investigated the timing of prebasic molt in adults of two migratory passerines, the endangered Black-capped Vireo (Vireo atricapilla) and the common White-eyed Vireo (V. griseus), on their breeding grounds in Texas. We compared the onset and rate of molt between sites differing in habitat and between years differing greatly in precipitation due to the El Nino–Southern Oscillation (ENSO). Black-capped Vireos began molting 13 days later (8 July) and completed molt 12 days faster (50 days) in low quality, clumped habitat than in more typical, contiguous shrubby habitat (62-day molt starting 25 June). Long-term estimates of onset and duration of molt based on study skins were intermediate. White-eyed Vireos in the site with clumped habitat a...

Increased Energy Expenditure but Decreased Stress Responsiveness during Molt

Baseline and stress-induced corticosterone (CORT), heart rate (fH), and energy expenditure were measured in eight captive European starlings Sturnus vulgaris during and following a prebasic molt. The fH and oxygen consumption (V O2 ) were measured simultaneously across a range of heart rates, and energy expenditure (kJ/d) was then calculated from data. Energy expenditure and fH were strongly and positively correlated in each individual. Baseline fH and energy expenditure were significantly higher during molt. Molting starlings expended 32% more energy over 24 h than nonmolting birds, with the most significant increase (60%) occurring at night, indicating a substantial energetic cost to molt. Furthermore, the cardiac and metabolic responses to stress during molt were different than during nonmolt. Birds were subjected to four different 30-min acute stressors. The fH and CORT responses to these stressors were generally lower during molt. Although restraint caused a 64% increase in daily energy expenditure during nonmolt, no other stressor caused a significant increase in energy expenditure. Overall, our data suggest that molt is not only energetically expensive but that it also alters multiple stress response pathways. Furthermore, most acute stressors do not appear to require a significant increase in energy expenditure.

Seasonal changes in adrenocortical responses to acute stress in Eurasian tree sparrow ( Passer montanus) on the Tibetan Plateau: Comparison with house sparrow ( P. domesticus) in North America and with the migratory P. domesticus in Qinghai Province

Seasonal modulation of the adrenocortical response to stress appears to be ubiquitous in arctic-breeding and temperate-zone-breeding birds, but has not been well investigated in alpine-breeding species at mid-latitude. We examined the adrenocortical response to acute stress (capture, handling and restraint) in populations of Eurasian tree sparrow ( Passer montanus) among seasons and migratory house sparrow ( P. domesticus bactrianus) in pre-breeding on the Qinghai-Xizang Plateau (the Tibetan Plateau). A population of house sparrow ( Passer domesticus domesticus) was also sampled in lowland Phoenix, Arizona during breeding and wintering stages. In Eurasian tree sparrows, baseline corticosterone (CORT) does not differ among life history stages, but stress-induced CORT level (maximal CORT, total and corrected integrated CORT) is significantly higher in late breeding stage than those in early breeding and prebasic molt stages. In house sparrows, stress-induced CORT level does not differ between sites and life history stages, but baseline CORT is significantly lower in pre-breeding from Qinghai compared with those in breeding and wintering stages from Phoenix. Interestingly, both baseline CORT and maximal CORT do not differ between the populations of Eurasian tree sparrow and house sparrow in early/pre-breeding stage although tree sparrow is resident species whereas house sparrow is migratory in Qinghai. Our results suggest that the extreme environment of the Tibetan Plateau does not have significant effects on adrenocortical responses to acute stress in Eurasian tree sparrows and house sparrows, which may be a result of masking by human activities. These invasive human commensals may have a unique HPA axis response to different environments because they can take advantage of human food sources and shelter (i.e. buildings).

Experimental manipulation of testosterone and condition during molt affects activity and vocalizations of male blue tits

Testosterone (T) is a key hormone regulating behavioral trade-offs in male birds, shifting investment towards sexual and competitive behaviors. However, the role of T in regulating male behavior during the molt has received very little attention, although this is a crucial life-history stage. Since the effect of T on behavior may be condition-dependent, particularly during the costly molt period, we studied the effects of T and condition in a two-way design. We manipulated T under two dietary regimes (standard and improved, resulting in an enhanced condition) in captive blue tits ( Cyanistes caeruleus) undergoing the first pre-basic molt. T treatment increased song frequency, indicating that song is T-dependent also at this time of year. Males on the improved diet sang less than males in relatively worse condition, providing no support for song as an indicator of male condition. T-treated males exhibited greater locomotor activity than control males, but only when fed the standard diet. Neither T- nor diet-treatment affected plumage maintenance (preening). Although T treatment resulted in a delay in molt progress all birds completed the molt. Taken together our results show that during the molt male birds are sensitive to relatively small fluctuations in T. Similar to its commonly observed effects during the breeding season, T stimulated an increase in song and locomotion. While there might be some benefits associated with such T effects, these must be traded-off against costs associated with conspicuous behavior and increased molt duration.

EFFECTS OF RAPID FLIGHT-FEATHER MOLT ON POSTBREEDING DISPERSAL IN A PURSUIT-DIVING SEABIRD

Abstract Breeding seabirds have been well studied but seabird ecology during the nonbreeding season is poorly understood because many species disperse far from breeding colonies to molt at sea. We characterized the timing of prebasic molt and postbreeding dispersal, described postbreeding dispersal movements, and estimated changes in body mass during molt for Marbled Murrelets (Brachyramphus marmoratus; Alcidae) in central California, 1999–2004. According to mark–recapture and at-sea surveys, 248–315 of 496–637 individuals (43–50%) used Ano Nuevo Bay, located immediately adjacent to nesting areas, for their prebasic molt in August–October. Long-distance dispersal (≥ 100 km) from Ano Nuevo Bay by radiomarked Marbled Murrelets was low during breeding (9–13%, n = 46), but was greater for individuals radiomarked at the end of the breeding season (69–90%, n = 20). The mean dispersal dates were 18 May and 21 October for the breeding and postbreeding samples, respectively, and postbreeding dispersal occurred an ...

Tracking Seasonal Movements of Adult Male Harlequin Ducks from Central Labrador Using Satellite Telemetry

Abstract Satellite telemetry was used to determine the migration patterns between breeding, staging, and molting areas of adult male Harlequin Ducks (Histrionicus histrionicus) breeding in central Labrador during 2001 (N = 5) and 2002 (N = 6). Male Harlequin Ducks remained near their capture locations for an average of 19-20 d. Males spent an average of 18 and 34 d on the Labrador coast in 2001 and 2002, respectively, and moved along the coast between several important staging areas prior to migration to Greenland. Male Harlequin Ducks may be completing a pre-basic molt prior to migrating to Greenland. Seven males migrated from the Labrador coast to three distinct molting areas in Greenland. All transmitters expired by September preventing a confirmation of whether these males wintered in Greenland. The absence of band sightings during winter in eastern Canada, especially Maine, indicates that some Harlequin Ducks breeding in central Labrador may also winter in eastern Greenland.

The effects of chronic psychological and physical stress on feather replacement in European starlings ( Sturnus vulgaris)

Corticosterone (CORT) is seasonally modulated in many passerines, with plasma CORT concentrations lowest during the prebasic molt, when all feathers are replaced. Recent evidence indicating that CORT implants slow the rate of feather regrowth in molting birds suggests that plasma CORT concentrations are downregulated during molt in order to avoid the inhibition of feather growth caused by the protein catabolic activity of CORT. To further test this hypothesis, we examined whether endogenous CORT release, stimulated by exposure to either psychological stress or physical stress (food restriction), could inhibit feather regrowth rates or decrease feather quality in birds undergoing an induced molt (feather replacement after plucking). European starlings ( Sturnus vulgaris) were exposed to chronic psychological stress or food restriction for three weeks of the feather regrowth period. Throughout this time, the length of growing primary, secondary, and tail feathers was measured and blood samples were collected to measure baseline and stress-induced CORT concentrations. Upon completion of growth, feather quality was analyzed via measurements of mass, rachis length, feather area, and presence of fault bars. Both psychological and physical stress protocols elevated circulating plasma CORT but significantly less than implants from an earlier study did. Psychological stress had no effect on feather regrowth rates or feather quality. Food restriction had no effect on feather growth rate but caused asynchronous feather replacement. When combined with psychological stress, physical stress also resulted in smaller feather area. Results indicate that CORT implants may not accurately represent chronic stress physiology. Additionally, the purpose for downregulating CORT concentrations during molt appears to be more complicated than simply protecting feather production from CORT's catabolic effects.

Repeatability of baseline corticosterone concentrations

One major assumption for endocrine studies is that hormone titers are consistent within an individual so that if hormone titers are low relative to the cohort on one day, they are relatively low compared to the cohort on other days. This is an especially important assumption for most field studies where researchers may have access to an individual animal only once. We used a laboratory study with captive house sparrows ( Passer domesticus) to test this assumption using glucocorticoid titers. Baseline corticosterone titers were measured five different times for each bird under six different experimental conditions: during both day and night while birds were held on a short day photoperiod (11L, 13D), a long day photoperiod (19L, 5D), and while birds were undergoing a prebasic molt. Although the variation within an individual was often larger than the variation between individuals, the relative ranks of birds compared to their cohort were consistent during the night in all three conditions. In contrast, during the day the relative ranks of birds compared to their cohorts were only consistent on short days; on long days and during molt there was no significantly consistent ranking among individuals. Furthermore, the overall rank of an individual in its cohort was often different during the day and night. These data indicate that it is not always a good assumption that birds can be categorized as individuals with higher and lower titers, which will complicate analyses of the causes of interindividual variation.

Prebasic molt initiation and progress in northern fulmars of the High Arctic: do molt and breeding overlap?

We examined feather molt progress of northern fulmars (Fulmarus glacialis) at Cape Vera in the Canadian High Arctic through opportunistic observation of individuals in flight from 2003 to 2006, and examination of bodies and wings of 127 individuals collected at the site, from 2003 to 2005. We found no evidence suggesting that fulmars shed primary feathers during breeding. Prebasic molt was initiated in the head, neck, sides, belly and back approximately 1 week before hatch. We failed to detect a sex effect on molt progress, but molt among breeders was delayed compared to molt in non- or failed breeders. This study constitutes a baseline we feel may be useful to: (1) researchers interested in feather replacement chronology, wherein feathers are used as sources of biological information; and (2) researchers interested in eventual assessment of relationships among large-scale environmental processes and molt progress in this species, especially in light of predicted changes to Arctic regions.

Rates Versus Counts: Fall Molts of Lucy's Warblers (Vermivora Luciae)

AbstractNew specimens from Sinaloa, Mexico, as well as two older specimens, show that both adult and recently fledged Lucy's Warblers (Vermivora luciae) often move far south of their breeding grounds to molt in the tropical deciduous forests of northwestern Mexico. Remarkably, the first preformative (= first prebasic) molt is complete in Lucy's Warblers, including the replacement of flight feathers, primary coverts, and rectrices; thus, previous aging criteria based on the wear and appearance of these feathers are invalid. We suggest that the recent conclusion that Lucy's Warblers molt on their breeding grounds (Voelker and McFarland 2002) is an artifact of more collecting in the southwestern United States than in western Mexico during late summer. An index of relative collecting in Mexico and the United States, as well as data from constant-effort mist netting, suggest that most Lucy's Warblers move south to molt.Tasas versus Conteos: Mudas de Otoño de Vermivora luciae

MOLT AND BODY MASS OF RED KNOTS IN THEEASTERN UNITED STATES

Red Knots (Calidris canutus) that spend winter in the southeastern United States are known to have been genetically separated from their congeners that migrate to Patagonian wintering grounds for about 12,000 years. We examined and documented differences between the two groups in their use of southward migration stopover locations, flight feather molt, fidelity to wintering zones, and differences in mass at southward migration stopover locations. Red Knots wintering in the southeastern United States do so consistently, and knots wintering in Patagonia have not changed to wintering in the southeastern United States. The two wintering groups have distinct differences in their nonbreeding season biology (e.g., migration strategies, chronology of pre-basic molt), and these differences have been maintained for decades if not millennia.

Photoperiodically-induced changes in hypothalamic–pituitary–adrenal axis sensitivity in captive house sparrows (Passer domesticus)

We used captive house sparrows (Passer domesticus) to identify regulatory mechanisms underlying seasonal (mimicked by changes in photoperiod) and diel differences in corticosterone output. Corticosterone responses were measured during three simulated seasons: short-day and long-day photoperiods and while birds underwent a pre-basic molt. Under all three conditions we tested for adrenal sensitivity by injecting exogenous ACTH, for pituitary sensitivity by injecting corticotropin-releasing factor (CRF) and arginine vasotocin (AVT), and for diel changes by repeating the injections during the day and at night. The daytime adrenal sensitivities were greatest on long days, lower on short days, and lowest during molt. These data suggest that reductions in either adrenal sensitivity to ACTH and/or capacity to secrete corticosterone could explain lowered endogenous corticosterone titers during molt. Furthermore, adrenal sensitivity to ACTH and pituitary sensitivity to AVT appeared to be greatest at night. This suggests that both the adrenal's sensitivity to the ACTH signal and the pituitary's capacity to secrete ACTH might provide a mechanism allowing for diel changes in corticosterone titers. This differs substantially from what is known about diel regulation in rodents. Taken together, these data provide further evidence that there are complex regulatory mechanisms controlling diel and seasonal changes in corticosterone titers in birds.

RATES VERSUS COUNTS: FALL MOLTS OF LUCY'S WARBLERS (VERMIVORA LUCIAE)

New specimens from Sinaloa, Mexico, as well as two older specimens, show that both adult and recently fl edged Lucy's Warblers (Vermivora luciae) oft en move far south of their breeding grounds to molt in the tropical deciduous forests of northwestern Mexico. Remarkably, the fi rst preformative (= fi rst prebasic) molt is complete in Lucy's Warblers, including the replacement of fl ight feathers, primary coverts, and rectrices; thus, previous aging criteria based on the wear and appear- ance of these feathers are invalid. We suggest that the recent conclusion that Lucy's Warblers molt on their breeding grounds (Voelker and McFarland 2002) is an artifact of more collecting in the southwestern United States than in western Mexico during late summer. An index of relative collecting in Mexico and the United States, as well as data from constant-eff ort mist nett ing, suggest that most Lucy's Warblers move south to molt. Received 11 November 2005, accepted 22 June 2006.