Interpopulation variation in contour feather structure is environmentally determined in great tits

Background

The plumage of birds is important for flying, insulation and social communication. Contour feathers cover most of the avian body and among other functions they provide a critical insulation layer against heat loss. Feather structure and composition are known to vary among individuals, which in turn determines variation in the insulation properties of the feather. However, the extent and the proximate mechanisms underlying this variation remain unexplored.

Methodology/Principal Findings

We analyzed contour feather structure from two different great tit populations adapted to different winter regimes, one northern population in Oulu (Finland) and one southern population in Lund (Sweden). Great tits from the two populations differed significantly in feather structure. Birds from the northern population had a denser plumage but consisting of shorter feathers with a smaller proportion containing plumulaceous barbs, compared with conspecifics from the southern population. However, differences disappeared when birds originating from the two populations were raised and moulted in identical conditions in a common-garden experiment located in Oulu, under ad libitum nutritional conditions. All birds raised in the aviaries, including adult foster parents moulting in the same captive conditions, developed a similar feather structure. These feathers were different from that of wild birds in Oulu but similar to wild birds in Lund, the latter moulting in more benign conditions than those of Oulu.

Conclusions/Significance

Wild populations exposed to different conditions develop contour feather differences either due to plastic responses or constraints. Environmental conditions, such as nutrient availability during feather growth play a crucial role in determining such differences in plumage structure among populations.     

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.     

The Extent of the Preserved Feathers on the Four-Winged Dinosaur Microraptor gui under Ultraviolet Light

Background

The holotype of the theropod non-avian dinosaur Microraptor gui from the Early Cretaceous of China shows extensive preservation of feathers in a halo around the body and with flight feathers associated with both the fore and hindlimbs. It has been questioned as to whether or not the feathers did extend into the halo to reach the body, or had disassociated and moved before preservation. This taxon has important implications for the origin of flight in birds and the possibility of a four-winged gliding phase.

Methodology/Principal Findings

Examination of the specimen under ultraviolet light reveals that these feathers actually reach the body of the animal and were not disassociated from the bones. Instead they may have been chemically altered by the body tissues of the animal meaning that they did not carbonise close into the animal or more likely were covered by other decaying tissue, though evidence of their presence remains.

Conclusions/Significance

These UV images show that the feathers preserved on the slab are genuinely associated with the skeleton and that their arrangement and orientation is likely correct. The methods used here to reveal hidden features of the specimen may be applicable to other specimens from the fossil beds of Liaoning that produced Microraptor.     

American Exceptionalism: Population Trends and Flight Initiation Distances in Birds from Three Continents

Background

All organisms may be affected by humans'' increasing impact on Earth, but there are many potential drivers of population trends and the relative importance of each remains largely unknown. The causes of spatial patterns in population trends and their relationship with animal responses to human proximity are even less known.

Methodology/Principal Finding

We investigated the relationship between population trends of 193 species of bird in North America, Australia and Europe and flight initiation distance (FID); the distance at which birds take flight when approached by a human. While there is an expected negative relationship between population trend and FID in Australia and Europe, we found the inverse relationship for North American birds; thus FID cannot be used as a universal predictor of vulnerability of birds. However, the analysis of the joint explanatory ability of multiple drivers (farmland breeding habitat, pole-most breeding latitude, migratory habit, FID) effects on population status replicated previously reported strong effects of farmland breeding habitat (an effect apparently driven mostly by European birds), as well as strong effects of FID, body size, migratory habit and continent. Farmland birds are generally declining.

Conclusions/Significance

Flight initiation distance is related to population trends in a way that differs among continents opening new research possibilities concerning the causes of geographic differences in patterns of anti-predator behavior.     

Allometry of the Duration of Flight Feather Molt in Birds

We used allometric scaling to explain why the regular replacement of the primary flight feathers requires disproportionately more time for large birds. Primary growth rate scales to mass (M) as M^0.171, whereas the summed length of the primaries scales almost twice as fast (M^0.316). The ratio of length (mm) to rate (mm/day), which would be the time needed to replace all the primaries one by one, increases as the 0.14 power of mass (M^0.316/M^0.171=M^0.145), illustrating why the time required to replace the primaries is so important to life history evolution in large birds. Smaller birds generally replace all their flight feathers annually, but larger birds that fly while renewing their primaries often extend the primary molt over two or more years. Most flying birds exhibit one of three fundamentally different modes of primary replacement, and the size distributions of birds associated with these replacement modes suggest that birds that replace their primaries in a single wave of molt cannot approach the size of the largest flying birds without first transitioning to a more complex mode of primary replacement. Finally, we propose two models that could account for the 1/6 power allometry between feather growth rate and body mass, both based on a length-to-surface relationship that transforms the linear, cylindrical growing region responsible for producing feather tissue into an essentially two-dimensional structure. These allometric relationships offer a general explanation for flight feather replacement requiring disproportionately more time for large birds.     

Haste makes waste: accelerated molt adversely affects the expression of melanin-based and depigmented plumage ornaments in house sparrows

Background

Many animals display colorful signals in their integument which convey information about the quality of their bearer. Theoretically, these ornaments incur differential production and/or maintenance costs that enforce their honesty. However, the proximate mechanisms of production costs are poorly understood and contentious in cases of non-carotenoid-based plumage ornaments like the melanin-based badge and depigmented white wing-bar in house sparrows Passer domesticus. Costly life-history events are adaptively separated in time, thus, when reproduction is extended, the time available for molt is curtailed and, in turn, molt rate is accelerated.

Methodology/Principal Findings

We experimentally accelerated the molt rate by shortening the photoperiod in order to test whether this environmental constraint is mirrored in the expression of plumage ornaments. Sparrows which had undergone an accelerated molt developed smaller badges and less bright wing-bars compared to conspecifics that molted at a natural rate being held at natural-like photoperiod. There was no difference in the brightness of the badge or the size of the wing-bar.

Conclusions/Significance

These results indicate that the time available for molt and thus the rate at which molt occurs may constrain the expression of melanin-based and depigmented plumage advertisements. This mechanism may lead to the evolution of honest signaling if the onset of molt is condition-dependent through the timing of and/or trade-off between breeding and molt.     

Solving a Migration Riddle Using Isoscapes: House Martins from a Dutch Village Winter over West Africa

Background

The ability to connect breeding, stopover and wintering locations of populations of migratory birds greatly enhances our understanding of the phenomenon of migration and improves our chances of effectively conserving these species. Among Palearctic-Afrotropical migratory species, aerial insectivores like the house martin (Delichon urbicum) are sensitive to factors influencing the availability of flying insects, and have declined in recent decades. The strict aerial behaviour of martins severely limits ring recoveries on wintering grounds and so there is a dearth of information on where European breeding populations over-winter in Africa, and the relative effects of population regulation on breeding vs. wintering grounds. We used a newly developed multi-isotope (δ ²H, δ ¹³C, δ ¹⁵N) feather isoscape for Africa together with inferences from summarized ring return data based on longitude, to assign winter origins to birds captured at a breeding colony in The Netherlands.

Principal Findings

Based on isotopic analyses of winter-grown martin feathers, we used a likelihood-based assignment approach to describe potential wintering locations where molt occurred of individual house martins from a Dutch colony by assigning them to four potential isotopically distinct clusters in Africa. We found the overwhelming majority of Dutch martins were assigned to a geographical cluster associated with West Africa.

Conclusions/Significance

The existence of strong isotopic gradients and patterns in African foodwebs that support migratory wildlife allows for the spatial assignment of tissues grown there. The assignment of Dutch house martins to wintering grounds primarily in West Africa was in strong agreement with independent and indirect methods used to infer winter origins of this species based on the association between the Normalized Difference Vegetation Index (NDVI) in Africa and population patterns in Italy and the United Kingdom. These confirmatory data-sets underscore the importance of suitable habitats in West Africa to the conservation of migratory aerial insectivores and other species.     

Plumage quality mediates a life-history trade-off in a migratory bird

Background

Moult is one of the most costly activities in the annual cycle of birds and most avian species separate moult from other energy-demanding activities, such as migration. To this end, young birds tend to undergo the first post-juvenile moult before the onset of migration, but in some species the time window for the pre-migratory feather replacement is too narrow. We hypothesized that in such species an increased investment in the structural quality of juvenile feathers may allow to retain juvenile plumage throughout the entire migratory period and delay moult until arriving at wintering grounds, thus avoiding a moult-migration overlap.

Methods

The effect of juvenile plumage quality on the occurrence of moult-migration overlap was studied in a migratory shorebird, the common snipe Gallinago gallinago. Ca. 400 of first-year common snipe were captured during their final stage of autumn migration through Central Europe. The quality of juvenile feathers was assessed as the mass-length residuals of retained juvenile rectrices. Condition of migrating birds was assessed with the mass of accumulated fat reserves and whole-blood hemoglobin concentration. Path analysis was used to disentangle complex interrelationships between plumage quality, moult and body condition.

Results

Snipe which grew higher-quality feathers in the pre-fledging period were less likely to initiate moult during migration. Individuals moulting during migration had lower fat loads and hemoglobin concentrations compared to non-moulting birds, suggesting a trade-off in resource allocation, where energetic costs of moult reduced both energy reserves available for migration and resources available for maintenance of high oxygen capacity of blood.

Conclusions

The results of this study indicate that a major life-history trade-off in a migratory bird may be mediated by the quality of juvenile plumage. This is consistent with a silver spoon effect, where early-life investment in feather quality affects future performance of birds during migration period. Our results strongly suggest that the juvenile plumage, although retained for a relatively short period of time, may have profound consequences for individuals’ fitness.

     

A Method for Investigating Population Declines of Migratory Birds Using Stable Isotopes: Origins of Harvested Lesser Scaup in North America

Background

Elucidating geographic locations from where migratory birds are recruited into adult breeding populations is a fundamental but largely elusive goal in conservation biology. This is especially true for species that breed in remote northern areas where field-based demographic assessments are logistically challenging.

Methodology/Findings

Here we used hydrogen isotopes (δD) to determine natal origins of migrating hatch-year lesser scaup (Aythya affinis) harvested by hunters in the United States from all North American flyways during the hunting seasons of 1999–2000 (n = 412) and 2000–2001 (n = 455). We combined geospatial, observational, and analytical data sources, including known scaup breeding range, δD values of feathers from juveniles at natal sites, models of δD for growing-season precipitation, and scaup band-recovery data to generate probabilistic natal origin landscapes for individual scaup. We then used Monte Carlo integration to model assignment uncertainty from among individual δD variance estimates from birds of known molt origin and also from band-return data summarized at the flyway level. We compared the distribution of scaup natal origin with the distribution of breeding population counts obtained from systematic long-term surveys.

Conclusions/Significance

Our analysis revealed that the proportion of young scaup produced in the northern (above 60°N) versus the southern boreal and Prairie-Parkland region was inversely related to the proportions of breeding adults using these regions, suggesting that despite having a higher relative abundance of breeding adults, the northern boreal region was less productive for scaup recruitment into the harvest than more southern biomes. Our approach for evaluating population declines of migratory birds (particularly game birds) synthesizes all available distributional data and exploits the advantages of intrinsic isotopic markers that link individuals to geography.     

A Quantitative Analysis of Flight Feather Replacement in the Moustached Tree Swift Hemiprocne mystacea,a Tropical Aerial Forager

The functional life span of feathers is always much less than the potential life span of birds, so feathers must be renewed regularly. But feather renewal entails important energetic, time and performance costs that must be integrated into the annual cycle. Across species the time required to replace flight feather increases disproportionately with body size, resulting in complex, multiple waves of feather replacement in the primaries of many large birds. We describe the rules of flight feather replacement for Hemiprocne mystacea, a small, 60g tree swift from the New Guinea region. This species breeds and molts in all months of the year, and flight feather molt occurs during breeding in some individuals. H. mystacea is one to be the smallest species for which stepwise replacement of the primaries and secondaries has been documented; yet, primary replacement is extremely slow in this aerial forager, requiring more than 300 days if molt is not interrupted. We used growth bands to show that primaries grow at an average rate of 2.86 mm/d. The 10 primaries are a single molt series, while the 11 secondaries and five rectrices are each broken into two molt series. In large birds stepwise replacement of the primaries serves to increase the rate of primary replacement while minimizing gaps in the wing. But stepwise replacement of the wing quills in H. mystacea proceeds so slowly that it may be a consequence of the ontogeny of stepwise molting, rather than an adaptation, because the average number of growing primaries is probably lower than 1.14 feathers per wing.     

Individual migratory patterns of two threatened seabirds revealed using stable isotope and geolocation analyses

Aim

Intraspecific variability in the migratory movements of seabirds is being revealed far more complex than hitherto recognized, and our lack of understanding undermines their effective protection. Our aim is to test whether the isotopic values of a single feather of two threatened seabirds, the Mediterranean (Puffinus yelkouan) and the Balearic (Puffinus mauretanicus) shearwaters allow the geographic assignment of birds to their non‐breeding areas.

Location

These two species are known to use three main non‐breeding areas: the NE Atlantic Ocean, the W Mediterranean and the Aegean‐Black seas.

Methods

We clustered in three groups the δ¹³C and δ¹⁵N values of the first primary feather (P1), inferred to be grown during the non‐breeding period, of 34 Mediterranean and 56 Balearic shearwaters accidentally caught by longliners. To link the isotopic values of P1 to its corresponding non‐breeding area, we performed a discriminant function analysis (DFA) based on the three clusters and applied this function to feathers of known origin: P1 from seven Mediterranean shearwaters from Hyères Archipelago (France) tracked with geolocators and body feathers from six chicks from Balearic shearwaters. To link the moulting patterns to the areas where the feathers were grown, we applied the DFA to a sequence of primary feathers of eight Balearic and five Mediterranean shearwaters (caught by longliners).

Results

Isotopic and tracking data indicate that none of the Mediterranean shearwaters migrated to the Atlantic. The cluster and discriminant function analyses revealed that 8% of Balearic and 54% of Mediterranean shearwater moulted P1 in the Mediterranean Sea. Migratory movements were reflected in the changing isotopic values of the primary sequences.

Main conclusions

Stable isotope analyses (SIA) are a powerful approach to reveal the intraspecific variability in the migratory patterns of seabirds that use distinct isotopic areas over their annual cycle. The assignment of birds to their non‐breeding areas by means of SIA is a simple and effective tool that can help to evaluate the impact of human activities in remote areas not only at population but also at individual level, which is an essential knowledge for the management and conservation of threatened species.

     

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

ABSTRACT 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 (δD_f) to 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 δD_f values of flight feathers grown at Arctic versus non‐Arctic latitudes enabled use of this technique to discriminate between age‐classes. We determined δD_f values of known Arctic‐grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather δD_f 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, δD_f 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.     

Flight modes in migrating European bee-eaters: heart rate may indicate low metabolic rate during soaring and gliding

Background

Many avian species soar and glide over land. Evidence from large birds (m _b>0.9 kg) suggests that soaring-gliding is considerably cheaper in terms of energy than flapping flight, and costs about two to three times the basal metabolic rate (BMR). Yet, soaring-gliding is considered unfavorable for small birds because migration speed in small birds during soaring-gliding is believed to be lower than that of flapping flight. Nevertheless, several small bird species routinely soar and glide.

Methodology/Principal Findings

To estimate the energetic cost of soaring-gliding flight in small birds, we measured heart beat frequencies of free-ranging migrating European bee-eaters (Merops apiaster, m _b∼55 g) using radio telemetry, and established the relationship between heart beat frequency and metabolic rate (by indirect calorimetry) in the laboratory. Heart beat frequency during sustained soaring-gliding was 2.2 to 2.5 times lower than during flapping flight, but similar to, and not significantly different from, that measured in resting birds. We estimated that soaring-gliding metabolic rate of European bee-eaters is about twice their basal metabolic rate (BMR), which is similar to the value estimated in the black-browed albatross Thalassarche (previously Diomedea) melanophrys, m _b∼4 kg). We found that soaring-gliding migration speed is not significantly different from flapping migration speed.

Conclusions/Significance

We found no evidence that soaring-gliding speed is slower than flapping flight in bee-eaters, contradicting earlier estimates that implied a migration speed penalty for using soaring-gliding rather than flapping flight. Moreover, we suggest that small birds soar and glide during migration, breeding, dispersal, and other stages in their annual cycle because it may entail a low energy cost of transport. We propose that the energy cost of soaring-gliding may be proportional to BMR regardless of bird size, as theoretically deduced by earlier studies.     

Linking hydrogen (δ2H) isotopes in feathers and precipitation: sources of variance and consequences for assignment to isoscapes

Background

Tracking small migrant organisms worldwide has been hampered by technological and recovery limitations and sampling bias inherent in exogenous markers. Naturally occurring stable isotopes of H (δ²H) in feathers provide an alternative intrinsic marker of animal origin due to the predictable spatial linkage to underlying hydrologically driven flow of H isotopes into foodwebs. This approach can assess the likelihood that a migrant animal originated from a given location(s) within a continent but requires a robust algorithm linking H isotopes in tissues of interest to an appropriate hydrological isotopic spatio-temporal pattern, such as weighted-annual rainfall. However, a number of factors contribute to or alter expected isotopic patterns in animals. We present results of an extensive investigation into taxonomic and environmental factors influencing feather δ ²H patterns across North America.

Principal Findings

Stable isotope data were measured from 544 feathers from 40 species and 140 known locations. For δ ²H, the most parsimonious model explaining 83% of the isotopic variance was found with amount-weighted growing-season precipitation δ ²H, foraging substrate and migratory strategy.

Conclusions/Significance

This extensive H isotopic analysis of known-origin feathers of songbirds in North America and elsewhere reconfirmed the strong coupling between tissue δ ²H and global hydrologic δ ²H patterns, and accounting for variance associated with foraging substrate and migratory strategy, can be used in conservation and research for the purpose of assigning birds and other species to their approximate origin.     

Does Feather Corticosterone Reflect Individual Quality or External Stress in Arctic-Nesting Migratory Birds?

The effects of environmental perturbations or stressors on individual states can be carried over to subsequent life stages and ultimately affect survival and reproduction. The concentration of corticosterone (CORT) in feathers is an integrated measure of hypothalamic–pituitary–adrenal activity during the molting period, providing information on the total baseline and stress-induced CORT secreted during the period of feather growth. Common eiders and greater snow geese replace all flight feathers once a year during the pre-basic molt, which occurs following breeding. Thus, CORT contained in feathers of pre-breeding individuals sampled in spring reflects the total CORT secreted during the previous molting event, which may provide insight into the magnitude or extent of stress experienced during this time period. We used data from multiple recaptures to disentangle the contribution of individual quality vs. external factors (i.e., breeding investment or environmental conditions) on feather CORT in arctic-nesting waterfowl. Our results revealed no repeatability of feather CORT within individuals of either species. In common eiders, feather CORT was not affected by prior reproductive investment, nor by pre-breeding (spring) body condition prior to the molting period. Individual feather CORT greatly varied according to the year, and August-September temperatures explained most of the annual variation in feather CORT. Understanding mechanisms that affect energetic costs and stress responses during molting will require further studies either using long-term data or experiments. Although our study period encompassed only five years, it nonetheless provides evidence that CORT measured in feathers likely reflects responses to environmental conditions experienced by birds during molt, and could be used as a metric to study carry-over effects.     

Interspecific comparison of the performance of soaring migrants in relation to morphology, meteorological conditions and migration strategies

Background

Performance of migrating birds can be affected by a number of intrinsic and extrinsic factors like morphology, meteorological conditions and migration strategies. We compared travel speeds of four raptor species during their crossing of the Sahara desert. Focusing the analyses on this region allows us to compare different species under equivalent conditions in order to disentangle which factors affect migratory performance.

Methodology/Principal Finding

We tracked raptors using GPS satellite transmitters from Sweden, Spain and Italy, and evaluated their migratory performance at both an hourly and a daily scale. Hourly data (flight speed and altitude for intervals of two hours) were analyzed in relation to time of day, species and season, and daily data (distance between roosting sites) in relation to species, season, day length and tailwind support.

Conclusions/Significance

Despite a clear variation in morphology, interspecific differences were generally very small, and did only arise in spring, with long-distance migrants (>5000 km: osprey and Western marsh-harrier) being faster than species that migrate shorter distances (Egyptian vulture and short-toed eagle). Our results suggest that the most important factor explaining hourly variation in flight speed is time of day, while at a daily scale, tailwind support is the most important factor explaining variation in daily distance, raising new questions about the consequences of possible future changes in worldwide wind patterns.     

Conserved syntenic clusters of protein coding genes are missing in birds

Background

Birds are one of the most highly successful and diverse groups of vertebrates, having evolved a number of distinct characteristics, including feathers and wings, a sturdy lightweight skeleton and unique respiratory and urinary/excretion systems. However, the genetic basis of these traits is poorly understood.

Results

Using comparative genomics based on extensive searches of 60 avian genomes, we have found that birds lack approximately 274 protein coding genes that are present in the genomes of most vertebrate lineages and are for the most part organized in conserved syntenic clusters in non-avian sauropsids and in humans. These genes are located in regions associated with chromosomal rearrangements, and are largely present in crocodiles, suggesting that their loss occurred subsequent to the split of dinosaurs/birds from crocodilians. Many of these genes are associated with lethality in rodents, human genetic disorders, or biological functions targeting various tissues. Functional enrichment analysis combined with orthogroup analysis and paralog searches revealed enrichments that were shared by non-avian species, present only in birds, or shared between all species.

Conclusions

Together these results provide a clearer definition of the genetic background of extant birds, extend the findings of previous studies on missing avian genes, and provide clues about molecular events that shaped avian evolution. They also have implications for fields that largely benefit from avian studies, including development, immune system, oncogenesis, and brain function and cognition. With regards to the missing genes, birds can be considered ‘natural knockouts’ that may become invaluable model organisms for several human diseases.

Electronic supplementary material

The online version of this article (doi:10.1186/s13059-014-0565-1) contains supplementary material, which is available to authorized users.     

Large-scale variation in flight feather molt as a mechanism enabling biennial breeding in albatrosses

Laysan and black-footed albatrosses, Phoebastria immutabilis and P . nigripes , exhibit both annual and biennial breeding frequencies, and annually replace flight feathers in patterns that can be described as large, small or medium in extent. Large molts are temporally incompatible with successful breeding. Small molts are temporally compatible with the longest breeding seasons. Medium molts are compatible with shorter, but still successful breeding seasons. On average, large and small molts combined replace the same feathers with the same frequencies as two medium molts combined. Thus, large and small annual molt patterns combined provide a mechanism for "transferring time from one year to another" enabling extended breeding seasons every other year, and thus biennial breeding. Medium-sized molts are compatible with annual breeding. Among multiple albatross species, large-scale, annual molt patterns can shift in response to shifting breeding frequencies, but there may be a time lag in the response. A newly identified period of rapid fattening following molt termination and preceding colony arrival suggests albatrosses maintain low fat stores throughout active molt to reduce wing-loading, intensifying temporal trade-offs between flight feather molt and breeding.     

Seasonal Changes in Colour: A Comparison of Structural,Melanin- and Carotenoid-Based Plumage Colours

Background

Plumage coloration is important for bird communication, most notably in sexual signalling. Colour is often considered a good quality indicator, and the expression of exaggerated colours may depend on individual condition during moult. After moult, plumage coloration has been deemed fixed due to the fact that feathers are dead structures. Still, many plumage colours change after moult, although whether this affects signalling has not been sufficiently assessed.

Methodology/Principal Findings

We studied changes in coloration after moult in four passerine birds (robin, Erithacus rubecula; blackbird, Turdus merula; blue tit, Cyanistes caeruleus; and great tit, Parus major) displaying various coloration types (melanin-, carotenoid-based and structural). Birds were caught regularly during three years to measure plumage reflectance. We used models of avian colour vision to derive two variables, one describing chromatic and the other achromatic variation over the year that can be compared in magnitude among different colour types. All studied plumage patches but one (yellow breast of the blue tit) showed significant chromatic changes over the year, although these were smaller than for a typical dynamic trait (bill colour). Overall, structural colours showed a reduction in relative reflectance at shorter wavelengths, carotenoid-based colours the opposite pattern, while no general pattern was found for melanin-based colours. Achromatic changes were also common, but there were no consistent patterns of change for the different types of colours.

Conclusions/Significance

Changes of plumage coloration independent of moult are probably widespread; they should be perceivable by birds and have the potential to affect colour signalling.