Differential expression of ASIP transcripts reveals genetic mechanism underpinning black-tail independence from body plumage in yellow-bodied chickens.

The genetic foundation of chicken body plumage color has been extensively studied. However, little attention has been paid to the inheritance patterns and molecular mechanisms underlying the formation of distal feather colors (tail and wingtip). Differences in these colors are common; for example, the Chinese Huiyang Beard chicken has black tail feathers, but yellow body plumage. Here, the hybrid offspring of Huiyang Beard and White Leghorn chickens were used to study the inheritance patterns of tail-feather color. The expression levels of pigment genes in differently colored feather follicles were analyzed using quantitative real-time PCR. The results showed that genetic regulation of tail-feather color was independent of body-plumage color. The Dominant White locus inhibited eumelanin synthesis in tail feathers without affecting the formation of yellow body plumage, whereas the Silver locus had the opposite effect. The expression of agouti signaling protein (ASIP) gene class 1 transcripts was significantly lower in black tail-feather follicles than in yellow body follicles, whereas tyrosinase-related protein 1 (TYRP1) gene expression was significantly higher in black tail feathers. These differentially expressed genes were confirmed to exert an effect on eumelanin and pheomelanin formation in feathers, thus influencing the regulation of chicken tail-feather color. In conclusion, this study lays the foundation for further research on the genetic mechanisms of regional differences in feather color, contributing to a better understanding of plumage pigmentation in chickens.

Plumage color degradation indicates reproductive effort: an experiment.

Plumage color has traditionally been regarded as a static ornamental trait, but evidence is accumulating for significant color changes without molt that typically reduce the conspicuousness of ornamentation. In some species, the social partner seems to increase its reproductive investment if the color trait is experimentally enhanced, suggesting that color change could act as a signal. However, the information content of this signal is so far unclear. For example, birds in poor condition or making greater effort may deteriorate more severely. We used brood size manipulations to alter the reproductive effort of male and female collared flycatchers Ficedula albicollis. Both sexes showed less severe decline in some reflectance attribute of their white breast when their brood was experimentally reduced. In each sex, greater deterioration of the reflectance trait affected by the manipulation was accompanied by increased feeding rate by the partner. These feeding patterns do not prove, but are consistent with, a compensatory response by the partner to induced degradation. The manipulation effects on color change we detected confirm for the first time that plumage color deterioration can indicate current reproductive effort, thereby providing a potential fitness advantage to social partners that react to such deterioration.

Urbanisation impacts plumage colouration in a songbird across Europe: Evidence from a correlational, experimental and meta-analytical approach.

Urbanisation is accelerating across the globe, transforming landscapes, presenting organisms with novel challenges, shaping phenotypes and impacting fitness. Urban individuals are claimed to have duller carotenoid-based colouration, compared to their non-urban counterparts, the so-called 'urban dullness' phenomenon. However, at the intraspecific level, this generalisation is surprisingly inconsistent and often based on comparisons of single urban/non-urban populations or studies from a limited geographical area. Here, we combine correlational, experimental and meta-analytical data on a common songbird, the great tit Parus major, to investigate carotenoid-based plumage colouration in urban and forest populations across Europe. We find that, as predicted, urban individuals are paler than forest individuals, although there are large population-specific differences in the magnitude of the urban-forest contrast in colouration. Using one focal region (Malmö, Sweden), we reveal population-specific processes behind plumage colouration differences, which are unlikely to be the result of genetic or early-life conditions, but instead a consequence of environmental factors acting after fledging. Finally, our meta-analysis indicates that the urban dullness phenomenon is well established in the literature, for great tits, with consistent changes in carotenoid-based plumage traits, particularly carotenoid chroma, in response to anthropogenic disturbances. Overall, our results provide evidence for uniformity in the 'urban dullness' phenomenon but also highlight that the magnitude of the effect on colouration depends on local urban characteristics. Future long-term replicated studies, covering a wider range of species and feeding guilds, will be essential to further our understanding of the eco-evolutionary implications of this phenomenon.

Covert-inspired flaps: an experimental study to understand the interactions between upperwing and underwing covert feathers.

Birds are agile flyers that can maintain flight at high angles of attack (AoA). Such maneuverability is partially enabled by the articulation of wing feathers. Coverts are one of the feather systems that has been observed to deploy simultaneously on both the upper and lower wing sides during flight. This study uses a feather-inspired flap system to investigate the effect of upper and lower side coverts on the aerodynamic forces and moments, as well as examine the interactions between both types of flaps. Results from wind tunnel experiments show that the covert-inspired flaps can modulate lift, drag, and pitching moment. Moreover, simultaneously deflecting covert-inspired flaps on the upper and lower sides of the airfoil exhibit larger force and moment modulation ranges compared to a single-sided flap alone. Data-driven models indicate significant interactions between the upper and lower side flaps, especially during the pre-stall regime for the lift and drag response. The findings from this study are also biologically relevant to the observations of covert feathers deployment during bird flight. Thus, the methods and results summarized here can be used to formulate new hypotheses about the coverts role in bird flight and develop a framework to design covert-inspired flow and flight control devices for engineered vehicles.

Integration of GWAS and eGWAS to screen candidate genes underlying green head traits in male ducks.

Sexually dimorphic plumage coloration is widespread in birds. The male possesses more brightly colored feathers than the female. Dark green head feathers comprise one of the most typical appearance characteristics of the male Ma duck compared with the female. However, there are noticeable individual differences observed in these characteristics. Herein, genome-wide association studies (GWAS) were employed to investigate the genetic basis of individual differences in male duck green head-related traits. Our results showed that 165 significant SNPs were associated with green head traits. Meanwhile, 71 candidate genes were detected near the significant SNPs, including four genes (CACNA1I, WDR59, GNAO1 and CACNA2D4) related to the individual differences in the green head traits of male ducks. Additionally, the eGWAS identified three SNPs located within two candidate genes (LOC101800026 and SYNPO2) associated with TYRP1 gene expression, and might be important regulators affecting the expression level of TYRP1 in the head skin of male ducks. Our data also suggested that transcription factor MXI1 might regulate the expression of TYRP1, thereby causing differences in the green head traits among male ducks. This study provided primary data for further analysis of the genetic regulation of duck feather color.

Plastic plumage colouration in response to experimental humidity supports Gloger's rule.

Knowing how animals adapt their phenotype to local temperature and humidity is key to understanding not only ecogeographical rules, but also how species will manage climate change, as current models predict changes in global patterns of temperature and precipitation. In endotherms, colour adaptations in response to climate have been under investigated, and their acclimatization-the individual capacity to reversibly adjust phenotype in response to different environments-is unknown. Geographic trends can provide clues about abiotic variables involved in colouration, as postulated by Gloger's rule, which predicts darker individuals in warm and humid regions. We tested whether house sparrows (Passer domesticus) can adjust colouration when faced with varying humidity conditions. We exposed birds to either a dry (humidity 45%) or a wet environment (70%) six months before their moult, and measured colouration in newly developed feathers in five parts of the body (bib, crown, crown stripe, belly and rump). As predicted by Gloger's rule, birds in wet conditions developed darker (bib and belly) and larger (bib) melanised plumage patches, than birds in dry conditions. Our result provides the first unequivocal evidence that the ability of individual birds to adjust their colouration may be a potential adaptation to climatic changes in endotherms.

Effect of polymorphisms in the 5'-flanking sequence of MC1R on feather color in Taihang chickens.

MC1R plays an important role in the regulation of the formation, transfer, and deposition of melanin in animals and is important for determining coat color. Many studies have reported on single nucleotide polymorphisms (SNPs) in the coding sequence of MC1R. However, few studies have investigated the polymorphisms in the 5'-flanking sequence of MC1R. In this study, we sequenced 2000 bp of the 5'-flanking sequence of MC1R in 300 Taihang chickens with brown feathers (MTH) and 300 Taihang chickens with black feathers (HTH). The sequencing results showed that 4 SNPs (MC1R g.18838722 G > C, g.18838624 T > C, g.18838694 G > A, and g.18838624 C > T) were located in the 5'-flanking sequence of MC1R between the MTH and HTH groups. Association analysis showed that there was a significant correlation between the 4 SNPs and feather color in Taihang chickens. The correlation between MC1R g.18838624 T >C and feather color of Taihang chicken was 100%, of which the CC (E1) genotype is MTH and the TT (E2) genotype is HTH. Furthermore, there was a significant correlation between MC1R g.18838624 T > C and egg production at 302 d. E1 (184.14 ± 0.674) was significantly higher than that in E2 (181.75 ± 0.577) (P < 0.05). Luciferase reporter assays were used to detect the transcriptional activity of MC1R with different SNP genotypes. The results showed that the luciferase activity of E2 was significantly higher than that of E1 (P < 0.05). In addition, transcription factor-binding site predictions showed that E2 creates a new binding site for ZEB1. RT‒qPCR results revealed that the expression of MC1R in E2 was significantly lower than that in E1 (P < 0.05), and the expression of ZEB1 in E2 was significantly higher than that in E1 (P < 0.05). Overexpression and shRNA experiments demonstrated that ZEB1 regulates the expression of MC1R in DF1 cells. ZEB1 has a negative regulatory effect on the transcriptional activity of MC1R; it inhibits the expression of MC1R and affects the feather color of Taihang chickens. This study provides new insight into the molecular mechanism of feather color formation and the transcriptional regulation of MC1R in Taihang chickens.

The evolutionary history and mechanistic basis of female ornamentation in a tropical songbird.

Ornamentation, such as the showy plumage of birds, is widespread among female vertebrates, yet the evolutionary pressures shaping female ornamentation remain uncertain. In part this is due to a poor understanding of the mechanistic route to ornamentation in females. To address this issue, we evaluated the evolutionary history of ornament expression in a tropical passerine bird, the White-shouldered Fairywren, whose females, but not males, strongly vary between populations in occurrence of ornamented black-and-white plumage. We first use phylogenomic analysis to demonstrate that female ornamentation is derived and that female ornamentation evolves independently of changes in male plumage. We then use exogenous testosterone in a field experiment to induce partial ornamentation in naturally unornamented females. By sequencing the transcriptome of experimentally induced ornamented and natural feathers, we identify genes expressed during ornament production and evaluate the degree to which female ornamentation in this system is associated with elevated testosterone, as is common in males. We reveal that some ornamentation in females is linked to testosterone and that sexes differ in ornament-linked gene expression. Lastly, using genomic outlier analysis we identify a candidate melanogenesis gene that lies in a region of high genomic divergence among populations that is also differentially expressed in feather follicles of different female plumages. Taken together, these findings are consistent with sex-specific selection favoring the evolution of female ornaments and demonstrate a key role for testosterone in generating population divergence in female ornamentation through gene regulation. More broadly, our work highlights similarities and differences in how ornamentation evolves in the sexes.

Practices and issues of moulting programs for laying hens: a review.

1. Moulting is a natural physiological process in birds when they shed their old feathers and replace them with new ones, and it is followed by reproductive quiescence resulting in reduced egg production. Different birds undergo moulting at different points in their life. Some birds have seasonal moulting while some moult at the end of their breeding cycle. This review will mainly focus on moulting practices associated with commercial layer birds because, in all other bird types, this is not managed.2. Commercial farms commonly analyse the cost-benefit ratio to decide the time and method to adopt for moulting. Commercial layer farms adopt different practices to force birds out of moult and restart the production cycle. Animal welfare groups consider this as stressful and against animal welfare, raising questions about the ethics of this practice.3. Many studies have been conducted using complete or partial feed withdrawal and non-feed withdrawal programs to measure their effectiveness in maintaining animal welfare, economy, and post-moult performance in mind.4. Animal welfare should not be compromised during moulting. The United States Egg Producers and other such groups from the United Kingdom and Europe have decided to sell eggs produced only through a non-feed withdrawal moulting programs.

Star finches Neochmia ruficauda have a visual preference for white dot patterns: a possible case of trypophilia.

Many animals have polka dot patterns on their body surface, some of which are known to have signalling functions; however, their evolutionary origins remain unclear. Dot patterns can trigger a fear response (trypophobia) in humans and are known to function as aposematic signals in non-human animals, suggesting that dots may deserve attention for biological reasons. Interestingly in many birds, plumage dot patterns serve for social/sexual signalling. To understand their evolution, we have focused on the sensory bias hypothesis, which predicts the role of pre-existing sensory preference driven by natural selection in shaping signal design. Our previous phylogenetic comparative study supported the hypothesis and showed that diet-driven visual preference promoted the evolution of plumage patterns, as there was an evolutionary correlation between termite-eating (white roundish gregarious prey) and the presence of plumage dot patterns in species of the family Estrildidae. This suggests that these species possess an intrinsic preference for dots. To test this, we compared the responses of an Estrildid species with dot plumage pattern (star finch Neochmia ruficauda) towards simultaneously presented monochrome-printed white dot vs white stripe patterns under both food-deprived and -supplied conditions. Overall, star finches preferred dots to stripes. They showed foraging-like behaviours almost only toward dots when hungry and gazed at dots frequently even when food was available, suggesting both hunger-related and hunger-neutral dot preferences. These results are rather surprising, given how strongly the subjects were attracted to abstract dot patterns without organic structure, but provided good support for the sensory bias hypothesis.

Effects of a FCBP gene polymorphism, location, and sex on Young's modulus of the tenth primary feather in racing pigeons.

Young's modulus (E) is a measure for stiffness of a material and a higher E means a higher stiffness. The respective polymorphism of the feather corneous beta-protein gene causes the replacement of glycine by cysteine. We looked for possible effects of the three FCBP genotypes on E in the 10th primaries of racing pigeons. However, we did not find a statistically significant difference of E between the genotypes, even within the sexes and/or within different locations under our test conditions. Our findings do not preclude the possibility that under other conditions (temperature, moisture) an influence of the glycine/cysteine polymorphism on E may exist. Compared to the more proximal locations of the rachis (base and middle) we observed lower values for E in the distal region (tip). The 10th primary constitutes the leading edge of the pigeon wing and this special function may require higher stiffness in the proximal parts of the shaft. We observed significantly higher values of E in females than in males, which result only from statistically significantly higher values in the middle region. The higher stiffness of female primaries may also contribute to the better results of hens compared to cocks in pigeon races.

Effects of stress during commercial hatching on growth, egg production and feather pecking in laying hens.

Every year, billions of egg layer chicks around the world are hatched under highly stressful, industrial circumstances. Here, it is investigated how the stressful procedure in the commercial hatchery, including incubation, hatching, processing, and transport affects the chicks with regards to traits relevant for the egg production industry. These traits were compared to those of a control group hatched in a small incubator and handled gently och quietly in a quiet room without any processing and transport. The chicks were weighed at hatch and at eight additional time points: 4 days, 1 week (w), 2 w, 3 w, 5 w, 8 w, 20 w and 25 w of age. Feather pecking was studied at 15 w of age and damages to the feathers and injuries on the comb and wattle were assessed at 25 w of age. From 19 w of age, eggs were collected on three days per week, counted and weighed. Chicks from a commercial hatchery had a lower hatch weight than control chicks (p<0.001). At 20 w of age, the weight of the commercial hatched chicks was still numerically lower, although this did not reach statistical significance. Commercially hatched chicks tended to show more feather pecking behaviour at 15 w of age compared to control chicks (p<0.1), although feather condition at 25 w of age showed the opposite pattern. Regarding production, commercially hatched chickens laid fewer (p<0.05) and smaller (p<0.05) eggs than chicks hatched and handled under calm circumstances. From this experiment, it is concluded that the stressful experience in the commercial hatchery has an overall negative effect on traits relevant for the industry.

Transcriptome profiling of morphogenetic differences between contour and flight feathers in duck.

1. This study examined the transcriptomic profiles of contour and flight feather follicles from two duck breeds to determine the molecular network and the candidate genes associated with contour and flight feather morphogenesis.2. High-throughput RNA sequencing was performed to compare differences in feather follicles between contour and flight feathers in two duck breeds (Heiwu and Nonghua duck).3. Comparing the contour feather follicles with flight feather follicles, 4,757 and 4,820 differentially expressed genes (DEGs) were identified in Heiwu and Nonghua duck respectively. Weighted gene co-expression network analysis (WGCNA) was used to construct a gene co-expression network of all DEGs and identify the key modules and hub genes associated with feather morphogenesis.4. Two key modules were enriched in many pathways involved in feather morphogenesis, such as the Wnt signalling pathway, anatomical structure morphogenesis, and focal adhesion. The CCNA2, TTK, NUF2, ECT2 and INCENP (in one module), and PRSS23, LAMC1, IGFBP3, SHISA5, and APLP2 (in another module) may be essential candidate genes for influencing feather morphology. Moreover, seven transcription factors (TFs) (UBP1, MBD2, ZNF512B, SMAD1, CAPN15, JDP2, KLF10, and MEF2A) were predicted to regulate the essential genes that contribute to feather morphogenesis.5. This work demonstrated gene expression changes of contour and flight feather follicles and is beneficial for further understanding of the complex structure of feathers.

Cis-acting mutation affecting GJA5 transcription is underlying the Melanotic within-feather pigmentation pattern in chickens.

Melanotic (Ml) is a mutation in chickens that extends black (eumelanin) pigmentation in normally brown or red (pheomelanin) areas, thus affecting multiple within-feather patterns [J. W. Moore, J. R. Smyth Jr, J. Hered. 62, 215-219 (1971)]. In the present study, linkage mapping using a back-cross between Dark Cornish (Ml/Ml) and Partridge Plymouth Rock (ml+/ml+ ) chickens assigned Ml to an 820-kb region on chromosome 1. Identity-by-descent mapping, via whole-genome sequencing and diagnostic tests using a diverse set of chickens, refined the localization to the genomic region harboring GJA5 encoding gap-junction protein 5 (alias connexin 40) previously associated with pigmentation patterns in zebrafish. An insertion/deletion polymorphism located in the vicinity of the GJA5 promoter region was identified as the candidate causal mutation. Four different GJA5 transcripts were found to be expressed in feather follicles and at least two showed differential expression between genotypes. The results showed that Melanotic constitutes a cis-acting regulatory mutation affecting GJA5 expression. A recent study established the melanocortin-1 receptor (MC1R) locus and the interaction between the MC1R receptor and its antagonist agouti-signaling protein as the primary mechanism underlying variation in within-feather pigmentation patterns in chickens. The present study advances understanding the mechanisms underlying variation in plumage color in birds because it demonstrates that the activity of connexin 40/GJA5 can modulate the periodic pigmentation patterns within individual feathers.

Exogenous corticosterone and melanin-based coloration explain variation in juvenile dispersal behaviour in the barn owl (Tyto alba).

Natal dispersal affects many processes such as population dynamics. So far, most studies have examined the intrinsic and extrinsic factors that determine the distance between the place of birth and of first breeding. In contrast, few researchers followed the first steps of dispersal soon after fledging. To study this gap, we radio-tracked 95 barn owl nestlings (Tyto alba) to locate their diurnal roost sites from the fledging stage until December. This was used to test whether the age of nest departure, post-fledging movements and dispersal distance were related to melanin-based coloration, which is correlated to fitness-related traits, as well as to corticosterone, a hormone that mediates a number of life history trade-offs and the physiological and behavioural responses to stressful situations. We found that the artificial administration of corticosterone delayed the age when juveniles left their parental home-range in females but not in males. During the first few months after fledging, longer dispersal distances were reached by females compared to males, by individuals marked with larger black feather spots compared to individuals with smaller spots, by larger individuals and by those experimentally treated with corticosterone. We conclude that the onset and magnitude of dispersal is sensitive to the stress hormone corticosterone, melanin-based coloration and body size.

Maternal age and maternal environment affect stress reactivity and measures of social behaviour in laying hens.

Maternal effects can shape the phenotypes of offspring, but the extent to which a layer breeder's experience can affect commercial laying hens remains unclear. We aimed to investigate the effects of maternal age and maternal environment on laying hens' behaviour and stress response. In our first experiment (E1), commercial hybrid hens were reared either in aviary or barren brooding cages, then housed in aviary, conventional cages or furnished (enriched) cages, thus forming different maternal housing treatments. Hens from each treatment were inseminated at three ages, and measures of response to manual restraint and social stress were assessed in offspring. In experiment 2 (E2), maternal age effects on offsprings' stress response were further investigated using fertile eggs from commercial breeder flocks at three ages. In E1, maternal age affected struggling and corticosterone during manual restraint, feather pecking and pulling and comb wounds. Additionally, maternal rearing and housing in aviary systems showed positive effects on measures of behaviour and stress response in offspring. Effects of maternal age were not replicated in E2, possibly due to methodological differences or higher tolerance to maternal effects in commercial breeders. Overall, we recommend researchers report parent stock age to increase comparison across studies and thus our understanding of maternal age effects.

Cyclic growth of dermal papilla and regeneration of follicular mesenchymal components during feather cycling.

How dermis maintains tissue homeostasis in cyclic growth and wounding is a fundamental unsolved question. Here, we study how dermal components of feather follicles undergo physiological (molting) and plucking injury-induced regeneration in chickens. Proliferation analyses reveal quiescent, transient-amplifying (TA) and long-term label-retaining dermal cell (LRDC) states. During the growth phase, LRDCs are activated to make new dermal components with distinct cellular flows. Dermal TA cells, enriched in the proximal follicle, generate both peripheral pulp, which extends distally to expand the epithelial-mesenchymal interactive interface for barb patterning, and central pulp, which provides nutrition. Entering the resting phase, LRDCs, accompanying collar bulge epidermal label-retaining cells, descend to the apical dermal papilla. In the next cycle, these apical dermal papilla LRDCs are re-activated to become new pulp progenitor TA cells. In the growth phase, lower dermal sheath can generate dermal papilla and pulp. Transcriptome analyses identify marker genes and highlight molecular signaling associated with dermal specification. We compare the cyclic topological changes with those of the hair follicle, a convergently evolved follicle configuration. This work presents a model for analyzing homeostasis and tissue remodeling of mesenchymal progenitors.

Age-dependent shifts and spatial variation in the diet of endangered Black-faced Spoonbill (Platalea minor) chicks.

The endangered Black-faced Spoonbill (Platalea minor) strictly breeds in marine environments and is threatened by the rapid loss of coastal wetlands within its breeding range. Adults with chicks are thought to gradually switch feeding sites from freshwater wetlands to coastal mudflats as the chicks' osmoregulatory system develops. We investigated age-dependent shifts in the diet of Black-faced Spoonbill chicks at four breeding colonies with varying freshwater habitat availability by examining stable isotopes (δ13C, δ15N) between the tip (grown at the age of 10 days) and middle (grown at the age of 22 days) portions of their primary feathers. The δ13C value of the middle portions was significantly higher than that of the tips, which suggested that the ratio of marine resources increased with the growth and development of chicks. A Bayesian isotope mixing model revealed that the diet proportion of marine prey in the early-chick rearing season was slightly higher than in the late-chick rearing season at three colonies in inshore areas, although this proportion was approximately 60% even in the early chick-rearing period. In contrast, isotopic values and reconstructed diet composition suggested that chicks in an offshore colony with limited freshwater wetlands relied more heavily on freshwater diets for both chick-rearing periods (>80%). Our results suggest that the shifts in feeding sites seen in previous studies might be related to the age-dependent dietary shift of chicks, highlighting the importance of freshwater wetlands for spoonbills on offshore islands without an inflow of freshwater in nearby intertidal mudflats. These findings emphasize the importance of freshwater prey and wetlands even for the endangered marine-breeding spoonbills, even though the negative impact of salt stress remains inconclusive.

Evolution of an Epidermal Differentiation Complex (EDC) Gene Family in Birds.

The transition of amniotes to a fully terrestrial lifestyle involved the adaptation of major molecular innovations to the epidermis, often in the form of epidermal appendages such as hair, scales and feathers. Feathers are diverse epidermal structures of birds, and their evolution has played a key role in the expansion of avian species to a wide range of lifestyles and habitats. As with other epidermal appendages, feather development is a complex process which involves many different genetic and protein elements. In mammals, many of the genetic elements involved in epidermal development are located at a specific genetic locus known as the epidermal differentiation complex (EDC). Studies have identified a homologous EDC locus in birds, which contains several genes expressed throughout epidermal and feather development. A family of avian EDC genes rich in aromatic amino acids that also contain MTF amino acid motifs (EDAAs/EDMTFs), that includes the previously reported histidine-rich or fast-protein (HRP/fp), an important marker in feather development, has expanded significantly in birds. Here, we characterize the EDAA gene family in birds and investigate the evolutionary history and possible functions of EDAA genes using phylogenetic and sequence analyses. We provide evidence that the EDAA gene family originated in an early archosaur ancestor, and has since expanded in birds, crocodiles and turtles, respectively. Furthermore, this study shows that the respective amino acid compositions of avian EDAAs are characteristic of structural functions associated with EDC genes and feather development. Finally, these results support the hypothesis that the genes of the EDC have evolved through tandem duplication and diversification, which has contributed to the evolution of the intricate avian epidermis and epidermal appendages.

Rapid adjustments of migration and life history in hemisphere-switching cliff swallows.

Many aspects of bird migration are necessarily innate.1 However, the extent of deterministic genetic control, environmental influence, and individual decision making in the control of migration remains unclear.2-8 Globally, few cases of rapid and dramatic life-history changes resulting in novel migration strategies are known. An example is latitudinal trans-hemispheric breeding colonization, whereby a subpopulation suddenly begins breeding on its non-breeding range.9-13 These life-history reversals demand concomitant changes in the timing of migration, feather molt, and breeding if the population is to remain viable.13 Cliff swallows, Petrochelidon pyrrhonota, are long-distance migrants that breed in North America and spend the non-breeding season mostly in South America.14 However, in 2015, a small population switched hemispheres by breeding successfully in Argentina,9 over 8,000 km from the nearest potential source, after presumably failed attempts.15,16 This provided a unique chance to characterize the early mechanisms of change in migratory behavior and phenology and to assess the possibility of double breeding. We tracked cliff swallows with geolocators following their second and fourth breeding seasons in Argentina, documenting inverted seasonality, three new migratory patterns and non-breeding areas (North America, Mesoamerica, and South America), and a shift of molt phenology by approximately 6 months, all possibly arising within a single generation. These birds did not practice migratory double breeding, although some spent the boreal summer in the traditional breeding range. Our data show that fundamental phenological changes occurred very rapidly during colonization and that phenotypic plasticity can underlie profound changes in the life histories of migratory birds.