Molting Birds Research Articles

Global review of quantitative studies of primary moult of birds using the Underhill-Zucchini moult model

Knowledge about the timing of moult and how it fits into the annual cycle, along with breeding and, in some cases migration, is an important component of understanding the life history of birds. In comparison with breeding and migration, there is a lack of knowledge about the moult of birds. Primary moult can be quantified using models such as the Underhill-Zucchini moult model. Studies that included estimates of primary moult parameters (duration, mean start date and standard deviation of mean start date) obtained using the Underhill-Zucchini moult model were compiled from the literature to generate a moult analysis database. These parameters provide information about the timing of moult, how it fits into the annual cycle and how synchronised moult is in the population. Relationships between the estimated moult parameters and a series of explanatory variables were explored. Critical gaps in our understanding of primary moult were revealed. The available data are taxonomically biased towards Charadriiformes and Passeriformes and there are substantial gaps in coverage across the phylogenetic tree. There are severe biases in the global distribution of moult studies across continents, with Africa having almost half of all applications of the moult model. Latitude emerged as an important explanatory variable but there are latitudinal gaps in the data, especially from the Tropic of Capricorn extending to 40°N and in the far northern and far southern hemispheres. Citizen scientists, both bird ringers and photographers, have a key role to play in helping to fill these data gaps. An emerging pattern in the available data is that the timing of primary moult is more synchronised in populations in far northern and far southern latitudes than in temperate or tropical zones, but more data in the far south is needed to confirm this. Timing of moult shows great variability in the tropics and sub-tropical regions of the southern hemisphere. In species that undertake an annual sequential moult, larger birds take longer to moult than smaller birds, but the allometric relationship is not tight. Migrants appear to be more synchronised in their moult than residents.

Physiological strategies of moult-migrating Black-necked Grebes (Podiceps nigricollis) in a polluted staging site according to blood chemistry

After breeding, Black-necked Grebes (Podiceps nigricollis) perform a moult-migration to autumn hypersaline staging sites, where they moult the flight feathers and forage on superabundant brine shrimp (Artemia spp.) before leaving for wintering areas. During the stay in moulting sites, the grebes experience changes in organs and muscle size (atrophy, hypertrophy), and almost double their body mass, which has been suggested to act as an insurance against a collapse in prey availability in late autumn. During two years we collected blood samples from hundreds of individuals at one of the most important European moulting sites (the Odiel marshes, SW Spain), which is a highly polluted area due to mining drainage and chemical industry. We assessed the potential effect of moulting stage, day of the year and body condition on 16 blood biochemical parameters. Because of the changes in prey availability and body composition of grebes, we expected some physiological adjustments during moult. Elevated levels of cholesterol suggested that birds in active moult increased foraging effort to face the costs of moulting. There was increased amount of lactate dehydrogenase, corresponding to periods of breast muscle atrophy. Birds in active moult augmented protein ingestion, likely to account for the requirements of feather growth. We also show that the probability of fasting due to low prey availability increased late in the moulting season, as demonstrated by an increase in plasma β-hydroxy-butyrate.

Book Review

Book Review

Transcriptomic Analysis of Laying Hens Revealed the Role of Aging-Related Genes during Forced Molting.

Molting in birds provides us with an ideal genetic model for understanding aging and rejuvenation since birds present younger characteristics for reproduction and appearance after molting. Forced molting (FM) by fasting in chickens causes aging of their reproductive system and then promotes cell redevelopment by providing water and feed again. To reveal the genetic mechanism of rejuvenation, we detected blood hormone indexes and gene expression levels in the hypothalamus and ovary of hens from five different periods during FM. Three hormones were identified as participating in FM. Furthermore, the variation trends of gene expression levels in the hypothalamus and ovary at five different stages were found to be basically similar using transcriptome analysis. Among them, 45 genes were found to regulate cell aging during fasting stress and 12 genes were found to promote cell development during the recovery period in the hypothalamus. In addition, five hub genes (INO80D, HELZ, AGO4, ROCK2, and RFX7) were identified by WGCNA. FM can restart the reproductive function of aged hens by regulating expression levels of genes associated with aging and development. Our study not only enriches the theoretical basis of FM but also provides insights for the study of antiaging in humans and the conception mechanism in elderly women.

Feather traits in four southern populations of the Eurasian blackcapSylvia atricapilla: do altitudinal movements explain the differences?

Moult of birds is shaped by environmental and genetic drivers whose relative contribution to the structure of feathers may differ within and between populations. In this study, we compare some traits of tail feathers (growth bars, mass, rachis width and barb length) between four populations of the Eurasian blackcapSylvia atricapillabreeding at different elevations within the southwestern Palaearctic. We tested if these traits were related to the primary productivity of habitats (a surrogate of food availability) or were better explained as an adaptation to altitudinal movements. The distribution of primary productivity was positively related to blackcap abundance suggesting that the species tracked the most productive areas to breed. In this environmental setting, wing morphology (wing length, concavity and pointedness) suggested that lowland blackcaps were sedentary while blackcaps from highland areas were involved in altitudinal movements. The feathers of blackcaps inhabiting the highlands showed wider growth bars and rachis than those of the most productive lowland areas but did not differ in feather mass and barb length. Fast feather growth has been related to time constraints to moult and wider rachis to improve flight efficiency in migratory birds. Our results therefore suggest that differences in feather characteristics between southern populations of the Eurasian blackcap are better interpreted as an adaptive response to altitudinal migration than as a consequence of regional food availability.

The Biology of Molt in Birds

The Biology of Molt in Birds

Seasonally divided moult in the Barred Warbler (Sylvia nisoria) is an endogenously controlled strategy

The seasonal timing of moult in migratory birds is an adaptation to cope with time constraints in the annual cycle. Kiat and Izhaki analysed moult patterns in Palaearctic passerines and rejected the proposition that seasonally divided moult is an endogenously controlled strategy. Instead, they advocated the view that it occurs due to a flexible and opportunistic timing of moult. In contrast, we argue that Kiat and Izhaki’s analysis is flawed and that they overlooked several important facts about moult in the Barred Warbler Sylvia nisoria and other species showing seasonally divided moult. These include the facts that juveniles replace a few secondaries already in their first winter, and that the moult sequences of primaries and secondaries are decoupled compared with the typical passerine complete moult sequence. We argue that seasonally divided moult is an adaptive strategy that is largely under endogenous control.

Marginal-covert moult in the House Sparrow Passer domesticus

ABSTRACT Marginal coverts are often neglected in passerine moult studies due to their small size and large number. Here, we document eight moult components of this group of wing coverts in a House Sparrow population from Barcelona in 2018–21: timing, growth rate, intensity, speed, duration, sequence, extent and barb density. We also tested the influence of various intrinsic and extrinsic factors on marginal-covert moult. Our adult population started marginal-covert moult on 29 July and finished it on 29 August, with the moult of hatch-year birds delayed 13 days in relation to the moult of adults. Growth rate of the largest marginal covert increased significantly with its size but did not differ between sexes. Intensity of marginal-covert moult replacement was not constant during the moult progress, which took 32 days on average. Moult sequence of marginal coverts started with the proximal feathers, proceeded distally replacing the central rows and was completed toward the front and rear rows. Moult extent was complete except in one case. Size of the largest marginal covert increased with calamus width, but not significantly, and barb density decreased with feather size, but not significantly, and neither measure differed between sexes. Comparison with published results indicated that growth rate is lower in marginal coverts than in primaries and rectrices, whereas barb density is greater. Linear regression of marginal-covert size on both growth rate and barb density can be used to predict growth rate and barb density of flight feathers in the House Sparrow.

"The Biology of Moult in Birds" by Lukas Jenni and Raffael Winkler, 2020. [book review

"The Biology of Moult in Birds" by Lukas Jenni and Raffael Winkler, 2020. [book review

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.

Local timing of rainfall predicts the timing of moult within a single locality and the progress of moult among localities that vary in the onset of the wet season in a year-round breeding tropical songbird

Rainfall seasonality is likely an important cue for timing key annual cycle events like moult in birds living in seasonally arid environments, but its precise effect is difficult to establish because seasonal rainfall may affect other covarying annual events such as breeding in the same way. In central Nigeria, however, Common Bulbuls Pycnonotus barbatus moult in the wet season but only show weak breeding seasonality. This suggests that moult is more sensitive to rainfall than breeding, but a similar outcome is possible if moult is simply periodic. We tested the relationship between rainfall and moult in Common Bulbuls at a single location over 18 years: on average moult started 5th May (± 41 days: 25th March–15th June), being on average later than the onset of the rains which is usually mid-April. The likelihood of finding a moulting Common bulbul was best predicted by rainfall 9–15 weeks before moult was scored. We then tested the generality of this across populations: the progress of moult should, therefore, correlate with the average timing of the wet season along a spatial environmental gradient where the rains start at different times each year south-to-north of Nigeria. To test this, we modelled moult progress just before the rains across 15 localities 6°–13° N as a function of the onset of the wet season among localities. As predicted, moult progressed further in localities with earlier wet seasons, confirming that the onset of moult is timed to the onset of the wet season in each locality despite weak breeding seasonality in the Common Bulbul. This strategy may evolve to maintain optimal annual cycle routine in seasonal environments where breeding is prone to unpredictable local perturbations like nest predation. It may, however, be less obvious in temperate systems where all annual cycle stages are seasonally constrained, but it may help with explaining the high frequency of breeding–moult overlaps in tropical birds.

Large‐scale longitudinal climate gradient across the Palearctic region affects passerine feather moult extent

Large‐scale spatial gradients of environmental conditions shape organisms, populations and ecosystems. Even though environmental gradients are a key research theme in macro‐ecology and biogeography, the effects of large‐scale, east–west, environmental gradients are largely overlooked compared with north–south gradients. Our study focused on feather moult, an important and energy demanding process in birds. By comparing Western and Eastern Palearctic populations of 21 species, we found that juvenile passerines in the Western and Eastern Palearctic differ in the number of feathers moulted as part of their post‐juvenile moult. This difference is most likely the result of a large‐scale climatic gradient in cold season duration and consequent differences in the time available for moulting. Eastern populations were characterized by a limited extent of feather moult that was additionally affected by migration distance and body mass. The longer migration distance in the Eastern Palearctic caused a generally less extensive moult while high body mass was correlated with a low difference in moult extent between the Western and Eastern Palearctic regions. These results highlight the importance of linking annual cycle processes at the organismal level to the specific environmental conditions within the distribution range of each species.

Factors controlling the migration phenology of Siberian Phylloscopus species

The timing of annual events in migratory species is determined by the position of breeding and wintering grounds, length of the breeding season, the occurrence of their preferred diet or the timing of moult in birds. Previous studies found significant differences in the migration phenology of Asian Phylloscopus warbler species, a group of long-distance migratory passerines. However, the factors that influence the observed differences in migration phenology have not been studied yet. Here, we analyze the effect of the above-mentioned factors on the migration timing of seven Phylloscopus species at a stopover site in the Russian Far East. We found that spring migration phenology was best explained by moult strategy and the preferred prey size, while autumn migration phenology was linked to the southernmost wintering latitude of the species. Interspecific differences in migration timing were much higher in autumn than in spring. The duration of the species-specific migration periods was also longer in autumn than in spring, most likely caused by higher competition during spring to arrive early at the breeding grounds. Our results contribute to the understanding of migration ecology in songbirds moving along the little-studied East Asian flyway.

First Report of Scoters (<i>Melanitta</i> spp.) along Eastern Baffin Island, Nunavut, Canada

In September of 2017 and 2018, we observed hundreds of scoters (Melanitta spp.) in fiords in Arctic Canada, approximately 1100 – 1800 km north of their previous northernmost observations. Given the remote locations and timing of observations, we do not know if these represent previously undiscovered areas where birds moult or new movements to Arctic locations. Moreover, the provenance of these sea ducks is unclear, as no evidence of movements to this region was indicated by large-scale satellite tracking of North American scoters during the last decade.

Post-Juvenile and Post-Breeding Moult of the Black Wheatear Oenanthe leucura in South-Eastern Spain: Slowed Down Due to Climatic Aridity?

Despite increasing interest in moult in birds, there are still clear gaps in our knowledge of this topic. For example, many aspects of the moult of several European passerines remain unknown. Here, I describe the extent of the postjuvenile and the post-breeding moult of Black Wheatears Oenanthe leucura in south-eastern Spain and analyse the possible influence of ecological factors on moult in this species. The post-juvenile moult of Black Wheatears involved a variable number of greater coverts (six in most cases), but tertials, alula feathers and the carpal covert were also moulted frequently. Primaries were never moulted, while secondaries and tail feathers were moulted in 1.5% of 65 birds. The number of feathers moulted in the post-juvenile moult was not related to sex or individuals' body size. The post-breeding moult occurred between late June and mid-November and followed the typical sequence of European passerines. It had a mean duration of 145 days, with no differences between sexes or years. The body condition of wheatears did not decrease as the moult advanced. The duration of the post-breeding moult of Black Wheatears was almost double the mean duration described for European passerines and is among the longest period ever described for passerines. The resident status and medium size of the species may partly explain the long duration of the post-breeding moult, but the low seasonality and high aridity of the study area seem to be the main drivers of the pattern found.

Determinants and constraints of feather growth.

During the periodic moult of the plumage of birds, a fast regrowth of feathers would shorten the time of reduced plumage functionality. However, it has long been known that feather growth-rate is limited and that long feathers take disproportionally longer to grow than small feathers, which has severe consequences on moult duration and the completeness of moult in large birds. The reasons for the limitations of feather-growth must be related to the size and/or functions of the feather follicle, but are largely unknown. Here we measured the size of the feather follicle (taking calamus width as a proxy) and related it to parameters of feather growth (feather growth-rate by mass and by length) and feather structure (feather length, mass, massiveness [mass of feather material per mm feather-length]). We used three independent datasets which allowed for interspecific analyses, and for intraspecific comparisons of differently structured feathers within the framework of biological scaling. We found that the cross-sectional area of the calamus (as a proxy of feather follicle size) was directly proportional to feather growth-rate by mass. Hence, factors acting at a two-dimensional scale (possibly nutrient supply to the growing feather) determines feather growth rate by mass, rather than the linear arrangement of stem cells (in a circular configuration) as had previously been assumed. Feather follicle size was correlated with both feather length and massiveness, hence it seems to be adapted to some extent to feather structure. Feather growth-rate by length was dependent on both the feather material produced per unit time (growth-rate by mass) and the amount of material deposited per unit feather-length. Follicle size not only determines feather growth-rate by mass, but also directly the structural design (shape, number of barbs, etc.) of a feather. Therefore, feather growth-rate is severely constrained by the requirements imposed by the structural feather design.

Speciation Associated with Shifts in Migratory Behavior in an Avian Radiation.

Migratory animals move up to thousands of kilometers every year [1]. Losses of migration (i.e., migratory drop-offs) occur when individuals of a migratory species stop migrating and establish founder sedentary populations, a phenomenon documented in birds [2-5] and butterflies [6]. In theory, losses-and also gains-of migration might promote speciation if sedentary and migratory populations become reproductively isolated [7-9]. Because migratory and sedentary strategies involve alternative physiological, behavioral, and morphological traits [10-13], divergence along multiple axes of organismal function is expected to accompany switches in migratory behavior, potentially accelerating speciation. We present evidence of speciation driven by a migratory drop-off in the fork-tailed flycatcher (Tyrannus savana) resulting in reproductive isolation likely driven by changes in breeding schedules (allochronic speciation [13-15]) and geographic isolation of breeding grounds (allopatric speciation [16]). Phylogenetic analyses across New World flycatchers (Tyrannidae) showed that an association between speciation and drop-offs is also observable at a macroevolutionary scale. Loss of migration was significantly more frequent than its gain, and speciation rates of migratory and partially migratory lineages (i.e., species having both migratory and sedentary populations) exceeded those of sedentary lineages. Models of trait evolution indicated that partial migration is an intermediate step between migratory and sedentary states in this family. Given that partial migration is widespread across migratory animals (e.g., of all migratory birds, ca. 51% are partially migratory [5]), speciation via switches in migratory behavior might be an important yet overlooked mechanism of animal diversification.

The Dynamics of the Replacement of Primary Flight Feathers in Passerine Birds and Possible Synchronization Factors

Abstract—A study was made to investigate the dynamics of the replacement of primary flight feathers in several passerine bird species in natural and laboratory conditions. Birds were kept at different photoperiods or early postbreeding molting was induced in winter by moving birds to a chamber with a 15-h photoperiod, which is efficient for gonadal development. Observations of birds were performed over 5 years for all photoperiods under study; a biorhythm with an approximately 3-day period was detected for the beginning of new flight feather development during molting in passerine birds. A time interval that was a multiple of approximately 3 days was observed between the replacements of adjacent primary flight features under various light–dark schedules (hours): 20 light (L) : 4 dark (D), 17L : 7D, 14L : 10D, and 12L : 12D. The phases of the 3-day biorhythm were compared with the dynamics of external atmospheric and heliogeophysical processes, and the most significant relationship was observed with extremes of changes in the speed of the Earth’s rotation around its axis. The external factors that are influenced by extremes of changes in the Earth’s rotational speed were assumed to act as synchronizers of the biorhythm.

Estimation of moult duration in birds with suspended moults: the case of the Red Crossbill and its relation to reproduction

Suspended moult is a relatively common phenomenon in birds, but it has remained relatively ignored for a long time in studies dealing with moulting process. Furthermore, the new and increasing number of models used to estimate moult duration systematically omit the fact that suspended moults can occur and that they significantly alter parameter estimates. Taking suspended moults into account is critical to obtain reliable moult-associated parameter estimates, which is fundamental given the demographic and evolutionary consequences of moult in birds. The main goal of this work is to develop a standardised method useful for characterising the main moulting parameters (such as the duration and starting date) of bird species that perform suspended moults. Additionally, with the aim of delving into our understanding of when and why suspended moults happen, we study their relationship with the occurrence of second breeding attempts in summer. We used data obtained from a Red Crossbill population in the Pyrenees during a period of 17 years. We observed that the percentage of crossbills undergoing suspended moult was approximately 50% during summer (July–August) and that moult suspension ultimately gives rise to a lengthened moulting period. Underhill–Zucchini models assume a time-linear replacement of feathers, overestimating the moult duration. Based on these same modelling approaches, we proposed a method to calculate the real moult duration after removing the time during which the moult was suspended. We also obtained evidence supporting the idea that crossbills suspend their moult as a strategy that could increase their breeding output in summer. The method proposed here can be used as a tool for working with species that undergo suspended moults.

Production and Well-Being Resulting From Delayed Movement of Pullets to the Hen Facility

SUMMARY Disease outbreaks can dramatically impact a producer's production schedule by disrupting the pullet movement into the hen house. One alternative is to early molt laying hens as they are coming into production. Early molted birds had lower production compared to controls with no differences in egg weight or egg components percentages. Bird welfare impacts were minimal as a result of the early molt. This management practice could be employed to allow pullets to be held minimizing production schedule disruptions as a result of a disease outbreak.