JOURNAL OF ORNITHOLOGY

Exploring host and geographical shifts in transmission of haemosporidians in a Palaearctic passerine wintering in India
Ishtiaq F
This is the first molecular study of avian haemosporidia diversity in wintering populations of the Blyth's Reed Warbler () in India that explores the extent of host and geographical shifts in transmission areas. In 156 birds, six lineages (37.8%; 95% CI 30.41-45.82%) and one lineage (1.9%; 95% CI 0.053-5.6%) were recovered. Of these, two lineages (ACDUM1 and ACDUM5) were detected in resident Himalayan birds, albeit in low frequency with absence of gametocytes in the blood suggesting no transmission from breeding to wintering quarters. In addition, there was no host sharing of local parasite lineages with the wintering populations.
Steroids in the Avian Brain: Heterogeneity across Space and Time
Schlinger BA
Sex steroids influence a diversity of neural and behavioral endpoints in birds, including some that have little to do with reproduction per se. Recent advances in neurochemistry and molecular biology further indicate that the avian brain is comprised of a network of unique sex steroid microenvironments. Factors involved in steroid synthesis and metabolism are present in the avian brain with expression levels that vary from region to region and with activities that are, in some cases, subject to regulation over relatively slow or rapid time intervals. Advances in our ability to a) isolate steroids from brain tissue and b) precisely measure their concentrations reveal how steroid levels vary spatially and temporally. A full appreciation of sex steroid effects on the avian brain require not only measures of hormones in blood but also an understanding of the numerous and varied mechanisms whereby the brain creates such a heterogeneous steroidal environment.
Unexpected dietary preferences of Eurasian Spoonbills in the Dutch Wadden Sea: spoonbills mainly feed on small fish not shrimp
Jouta J, de Goeij P, Lok T, Velilla E, Camphuysen CJ, Leopold M, van der Veer HW, Olff H, Overdijk O and Piersma T
After an historical absence, over the last decades Eurasian Spoonbills have returned to breed on the barrier islands of the Wadden Sea. The area offers an abundance of predator-free nesting habitat, low degrees of disturbance, and an extensive intertidal feeding area with increasing stocks of brown shrimp , the assumed main prey of . Nevertheless, newly established and expanding colonies of spoonbills have surprisingly quickly reached plateau levels. Here we tested the often stated assertion that spoonbills mainly rely on brown shrimp as food, by quantifying the diet of chicks on the basis of regurgitates and by analysis of blood isotopes using stable isotope Bayesian mixing models. Both methods showed that, rather than brown shrimp being the staple food of spoonbill chicks, small flatfish (especially plaice ) and gobies ( spp.) were their main prey. Unlike shrimp, small flatfish have been reported to be rather scarce in the Wadden Sea in recent years, which may explain the rapid saturation of colony size due to food-related density-dependent recruitment declines of growing colonies. By way of their diet and colony growth characteristics, spoonbills may thus indicate the availability of small fish in the Wadden Sea. We predict that the recovery to former densities of young flatfish and other juvenile/small fish in the Wadden Sea will be tracked by changing diets (more fish) and an increase in the size of Eurasian Spoonbill colonies across the Wadden Sea.
The impact of invasive plant management on the foraging ecology of the Warbler Finch () and the Small Tree Finch () on Galápagos
Filek N, Cimadom A, Schulze CH, Jäger H and Tebbich S
In recent decades, arboreal Darwin's Finches have suffered from a dramatic population decline, which has been attributed to parasitism by the invasive botfly . However, changes to their primary habitat caused by invasive plant species may have additionally contributed to the observed population decline. The humid cloud forest on Santa Cruz Island is a stronghold of arboreal Darwin's Finches but has been invaded by blackberry (). In some areas, manual control and herbicide application are used to combat this invasion, both causing a temporary removal of the entire understory. We hypothesized that the removal of the understory reduces the availability of arthropods, which are a main food source during chick rearing. We compared the foraging behaviour of Warbler Finches () and Small Tree Finches () at three study sites that varied in the degree of invasion and the length of time since the last herbicide application. We used prey attack rate and foraging success as an index for food availability and predicted a lower attack rate and foraging success in areas that had recently been sprayed with herbicides. We found that both the invasion and the management of influenced microhabitat use, foraging substrate and prey choice in both species. Contrary to our hypothesis, we did not find a lower attack rate or foraging success in the area with recent herbicide application. This may be explained by the finding that both species mainly foraged in the canopy but also used dead plant structures of the understory of the recently controlled area that resulted from the invasive plant management.
Parental behaviour and family proximity as key to gosling survival in Greylag Geese ()
Szipl G, Loth A, Wascher CAF, Hemetsberger J, Kotrschal K and Frigerio D
Reproductive success in monogamous species is generally affected by both behavioural and hormonal fine-tuning between pair partners. Vigilance, defence and brooding of offspring are among the main parental investments, and often the sexes adopt different roles. In the present study, we investigate how sex differences in parental behaviour and family proximity in the socially monogamous Greylag Goose () affect gosling survival. During the reproductive season in spring 2013, we recorded the behaviour of 18 pairs with offspring and gosling survival in a semi-tame, long-term monitored, and individually marked flock of Greylag Geese in Grünau, Austria. We found that behavioural role differentiation between the parents varied with developmental phase, and thus with gosling age. Especially during the first 10 days after hatching, females were foraging more frequently than males, which were more vigilant and aggressive towards other flock members. Such differences between the sexes levelled out 20 to 30 days after hatching. In general, females stayed in closer proximity to their offspring than males. Gosling survival was high when the parents were relatively aggressive and emphasized vigilance rather than foraging behaviour. Hence, we show a direct link between pair partners' quality of parental investment and gosling survival.
Age effects in Darwin's finches: older males build more concealed nests in areas with more heterospecific singing neighbors
Huge AC, Adreani NM, Colombelli-Négrel D, Akçay Ç, Common LK and Kleindorfer S
Nesting success tends to increase with age in birds, in part because older birds select more concealed nest sites based on experience and/or an assessment of prevailing predation risk. In general, greater plant diversity is associated with more biodiversity and more vegetation cover. Here, we ask if older Darwin's finch males nest in areas with greater vegetation cover and if these nest sites also have greater avian species diversity assessed using song. We compared patterns in Darwin's Small Tree Finch () and Darwin's Small Ground Finch () as males build the nest in both systems. We measured vegetation cover, nesting height, and con- vs. heterospecific songs per minute at 55 nests (22 , 33 ). As expected, in both species, older males built nests in areas with more vegetation cover and these nests had less predation. A novel finding is that nests of older males also had more heterospecific singing neighbors. Future research could test whether older males outcompete younger males for access to preferred nest sites that are more concealed and sustain a greater local biodiversity. The findings also raise questions about the ontogenetic and fitness consequences of different acoustical experiences for developing nestlings inside the nest.
Migration and non-breeding ecology of the Yellow-breasted Chat
Mancuso KA, Hodges KE, Alexander JD, Grosselet M, Bezener AM, Morales L, Martinez SC, Castellanos-Labarcena J, Russello MA, Rockwell SM, Bieber ME and Bishop CA
Detailed information spanning the full annual cycle is lacking for most songbird populations. We examined breeding, migration, and non-breeding sites for the Yellow-breasted Chat ( chat). We deployed archival GPS tags and light-level geolocators on breeding chats in British Columbia and light-level geolocators in California from 2013 to 2017 to determine migration routes and non-breeding sites. We examined whether chats overwintered in protected areas and characterized the percent of land cover within 1 km. We used a combination of genetics and stable hydrogen isotopes from feathers collected on non-breeding chats in Nayarit, Mexico (2017-2019) and migrating chats in Chiapas, Mexico (2018) and Veracruz, Mexico (2014-2015) to determine subspecies and infer breeding location. Endangered chats in British Columbia followed the Pacific Flyway and spent the non-breeding period in Sinaloa and Nayarit, Mexico. Two out of five chats spent the non-breeding period in protected areas, and the most common landcover type used was tropical or subtropical broadleaf deciduous forest. We found no mixing of eastern and western chats in our Mexico sites, suggesting strong migratory connectivity at the subspecies level. Western chats likely originating from multiple breeding latitudes spent the non-breeding period in Nayarit. Eastern Yellow-breasted Chats likely breeding across various latitudes migrated through Veracruz and Chiapas. Our results provide precise migration routes and non-breeding locations, and describe habitat cover types for chats, notably an endangered population in British Columbia, which may be valuable for habitat protection and conservation efforts.
Insights into the migration of the European Roller from ring recoveries
Finch T, Dunning J, Kiss O, Račinskis E, Schwartz T, Sniauksta L, Szekeres O, Tokody B, Franco A and Butler SJ
Despite recent advances in avian tracking technology, archival devices still present several limitations. Traditional ring recoveries provide a complementary method for studying migratory movements, particularly for cohorts of birds with a low return rate to the breeding site. Here we provide the first international analysis of ring recovery data in the European Roller , a long-distance migrant of conservation concern. Our data comprise 58 records of Rollers ringed during the breeding season and recovered during the non-breeding season. Most records come from Eastern Europe, half are of juveniles and over three quarters are of dead birds. Thus, ring recoveries provide migration data for cohorts of Rollers-juveniles and unsuccessful migrants-for which no information currently exists, complementing recent tracking studies. Qualitatively, our results are consistent with direct tracking studies, illustrating a broad-front migration across the Mediterranean Basin in autumn and the use of the Arabian Peninsula by Rollers from eastern populations in spring. Autumn movements were, on average, in a more southerly direction for juveniles than adults, which were more easterly. Juvenile autumn recovery direction also appeared to be more variable than in adults, though this difference was not statistically significant. This is consistent with juveniles following a naïve vector-based orientation program, and perhaps explains the 'moderate' migratory connectivity previously described for the Roller. In the first (qualitative) analysis of Roller non-breeding season mortality, we highlight the high prevalence of shooting. The recovery age ratio was juvenile-biased in autumn but adult-biased in spring. Although not statistically significant, this difference points towards a higher non-breeding season mortality of juveniles than adults. Our study demonstrates the complementarity of ring recoveries to direct tracking, providing an insight into the migration of juvenile Rollers and non-breeding season mortality.
Community engagement and data quality: best practices and lessons learned from a citizen science project on birdsong
Jäckel D, Mortega KG, Darwin S, Brockmeyer U, Sturm U, Lasseck M, Moczek N, Lehmann GUC and Voigt-Heucke SL
Citizen Science (CS) is a research approach that has become popular in recent years and offers innovative potential for dialect research in ornithology. As the scepticism about CS data is still widespread, we analysed the development of a 3-year CS project based on the song of the Common Nightingale () to share best practices and lessons learned. We focused on the data scope, individual engagement, spatial distribution and species misidentifications from recordings generated before (2018, 2019) and during the COVID-19 outbreak (2020) with a smartphone using the 'Naturblick' app. The number of nightingale song recordings and individual engagement increased steadily and peaked in the season during the pandemic. 13,991 nightingale song recordings were generated by anonymous (64%) and non-anonymous participants (36%). As the project developed, the spatial distribution of recordings expanded (from Berlin based to nationwide). The rates of species misidentifications were low, decreased in the course of the project (10-1%) and were mainly affected by vocal similarities with other bird species. This study further showed that community engagement and data quality were not directly affected by dissemination activities, but that the former was influenced by external factors and the latter benefited from the app. We conclude that CS projects using smartphone apps with an integrated pattern recognition algorithm are well suited to support bioacoustic research in ornithology. Based on our findings, we recommend setting up CS projects over the long term to build an engaged community which generates high data quality for robust scientific conclusions.
A low-cost wind tunnel for bird flight experiments
Grogger HA, Gossar M, Makovec M, Fritz J, Neugebauer K, Amann F and Voelkl B
A blower-type wind tunnel for physiological bird flight experiments has been developed, constructed and evaluated. Since the birds to be investigated are rather big (Northern Bald Ibis, ), the cross-sectional area of the test section measures 2.5 m × 1.5 m. The maximum achievable flow speed is approximately 16 ms. The wind tunnel exhibits a flexible outlet nozzle to provide up- and downdraft to allow for gliding and climbing flights. The current paper describes in detail the layout, design and construction of the wind tunnel including its control. Numerical simulations of the flow and measurements of the velocity distribution in the test section are presented. Apart from a non-homogeneous flow region in the mixing layer at the boundaries of the free jet, the test section exhibits a very even velocity distribution; the local speed deviates by less than two percent from the mean velocity. The turbulence intensity inside the test section was measured to be between 1 and 2%. As a constraint, a limited budget was available for the project. Four northern bald ibises were hand-raised and trained to fly in the wind tunnel.
Parental food provisioning and nestling growth under parasitism in the Galapagos Green Warbler-Finch, classified as 'vulnerable' by the IUCN
Pike CL, Kofler B, Richner H and Tebbich S
In the Galapagos Islands, many endemic landbird populations are declining due to habitat degradation, food availability, introduced species and other factors. Given nestlings typically lack efficient defense mechanisms against parasites, hematophagous ectoparasites such as the larvae of the introduced Avian Vampire Fly, , can impose high brood mortality and cause threatening population declines in Darwin finches and other landbirds. Here, we assess whether the food compensation hypothesis (i.e., the parents' potential to compensate for deleterious parasite effects via increased food provisioning) applies to the Green Warbler-Finch. We differentiated nests with low or high infestation levels by . and quantified food provisioning rates of male and female parents, time females spent brooding nestlings, and nestling growth. Male provisioning rates, total provisioning rates and female brooding time did not significantly vary in relation to infestation levels, nor by the number of nestlings. Opposed to the predictions of the food compensation hypothesis, females showed significantly reduced provisioning rates at high infestation levels. Nestling body mass was significantly lower and there was a reduction of skeletal growth, although not significantly, in highly infested nests. The females' response to high infestation may be due to parasites directly attacking and weakening brooding females, or else that females actively reduce current reproductive effort in favor of future reproduction. This life-history trade-off may be typical for Darwin finches and many tropical birds with long lifespans and therefore high residual reproductive value. Conservation strategies may not build on the potential for parental food compensation by this species.
Kea, bird of versatility. Kea parrots () show high behavioural flexibility in solving a demonstrated sequence task
Suwandschieff E, Huber L, Bugnyar T and Schwing R
Social learning is an important aspect of dealing with the complexity of life. The transmission of information via the observation of other individuals is a cost-effective way of acquiring information. It is widespread within the animal kingdom but may differ strongly in the social learning mechanisms applied by the divergent species. Here we tested eighteen Kea () parrots on their propensity to socially learn, and imitate, a demonstrated sequence of steps necessary to open an apparatus containing food. The demonstration by a conspecific led to more successful openings by observer birds, than control birds without a demonstration. However, all successful individuals showed great variation in their response topography and abandoned faithfully copying the task in favour of exploration. While the results provide little evidence for motor imitation they do provide further evidence for kea's propensity towards exploration and rapidly shifting solving strategies, indicative of behavioural flexibility.
Why are ravens smart? Exploring the social intelligence hypothesis
Bugnyar T
Ravens and other corvids are renowned for their 'intelligence'. For long, this reputation has been based primarily on anecdotes but in the last decades experimental evidence for impressive cognitive skills has accumulated within and across species. While we begin to understand the building blocks of corvid cognition, the question remains why these birds have evolved such skills. Focusing on Northern Ravens , I here try to tackle this question by relating current hypotheses on brain evolution to recent empirical data on challenges faced in the birds' daily life. Results show that foraging ravens meet several assumptions for applying social intelligence: (1) they meet repeatedly at foraging sites, albeit individuals have different site preferences and vary in grouping dynamics; (1) foraging groups are structured by dominance rank hierarchies and social bonds; (3) individual ravens memorize former group members and their relationship valence over years, deduce third-party relationships and use their social knowledge in daily life by supporting others in conflicts and intervening in others' affiliations. Hence, ravens' socio-cognitive skills may be strongly shaped by the 'complex' social environment experienced as non-breeders.
Temperature and aridity determine body size conformity to Bergmann's rule independent of latitudinal differences in a tropical environment
Nwaogu CJ, Tieleman BI, Bitrus K and Cresswell W
Bergmann's rule, defined as the tendency for endotherms to be larger in colder environments, is a biophysical generalization of body size variation that is frequently tested along latitudinal gradients, even though latitude is only a proxy for temperature variation. We test whether variation in temperature and aridity determine avian body size conformity to Bergmann's rule independent of latitude differences, using the ubiquitous Common Bulbul , along a West African environmental gradient. We trapped 538 birds in 22 locations between latitudes 6 and 13°N in Nigeria, and estimated average body surface area to mass ratio per location. We then modelled body surface to mass ratio using general linear models, with latitude, altitude and one of 19 bioclimatic variables extracted from http://www.worldclim.org/bioclim as predictors. We sequentially dropped latitude and altitude from each model to obtain the of the resultant models. Finally, we compared the of univariate models, where bioclimatic variables predicted body surface area to mass ratio significantly (14 out of 19), to multivariate models including latitude, altitude and a bioclimatic variable, using the Wilcoxon matched pairs test. We found that multivariate models did not perform better than univariate models with only bioclimatic variables. Six temperature and eight precipitation variables significantly predicted variation in body surface area to mass ratio between locations; in fact, 50% (seven out of 14) of these better explained variation in body surface area to mass ratio than the multivariate models. Birds showed a larger body surface area relative to body mass ratio in hotter environments independent of latitude or altitude, which conforms to Bergmann's rule. Yet, a combination of morphometric analyses and controlled temperature-exposure experiments is required to prove the proposed relationship between relative body surface area and thermoregulation in endotherms.
No evidence that nest site choice in Pied Flycatchers is mediated by assessing the clutch size of a heterospecific, the Great Tit
Slagsvold T and Wiebe KL
Among species that use similar resources, an individual may benefit by observing and copying the behavioural decision of a heterospecific. We tested the hypothesis of heterospecific social learning in passerine birds, namely that a migrant species, the Pied Flycatcher uses external markings on the nest cavities of a resident species, the Great Tit , as cues when choosing a nest site. Others have suggested that prospecting flycatchers assess the clutch size of tit "demonstrators" by entering their nest boxes and, assuming that a large clutch indicates a high-quality individual, will copy the nest appearance of tits with large, but not small clutches. During a 4-year period in Norway, we designed a similar study but did not find that flycatchers based their nest choice on the clutch size of tits. Neither were there any relationships between clutch size of the tit and its laying date, incubation behaviour, or the number of eggs visible through nest material during egg-laying so Pied Flycatchers did not use these indirect cues to assess quality of the tutor. Filming of tit nests showed that prospecting flycatchers did not enter tit nest boxes to assess the content. Indeed, incubating female tits only left their nest boxes for short bouts of unpredictable duration so there was little opportunity for flycatchers to inspect the nest contents unnoticed. Our study calls into question the mechanism of using the content of tit nests as public information for choosing traits of nest sites based on external characteristics. We suggest that similar studies of nest site choice in relation to possible social information transfer be replicated more widely.
Fecal sacs do not increase nest predation in a ground nester
Rubio E, Sanllorente O, Tieleman BI and Ibáñez-Álamo JD
Most altricial birds remove their nestlings' feces from the nest, but the evolutionary forces driving this behavior are poorly understood. A possible adaptive explanation for this could be that birds avoid the attraction of nest predators to their nests due to the visual or olfactory cues produced by feces (nest predation hypothesis). This hypothesis has received contrasting support indicating that additional experimental studies are needed, particularly with respect to the visual component of fecal sacs. To test this hypothesis, we conducted an experiment manipulating the presence of fecal sacs on inactive Woodlark () nests. This ground nester has highly cryptic nests that are mainly depredated by visually oriented nest predators (i.e., corvids) in our study population, making it an excellent system to test for the nest predation hypothesis. Our results showed that the presence of fecal sacs in the nest does not seem to be an important factor explaining nest predation. Interestingly, the effect of nest concealment, the most important factor explaining nest predation in Woodlark nests, depended on whether the nest was depredated the previous year or not, supporting the importance of using different nesting sites between years. Our findings indicate that this important nest sanitation behavior is not likely motivated by nest predation and highlight the need to explore alternative selective pressures in this context.
Moult of overwintering Wood Warblers in an annual-cycle perspective
Jarrett C, Powell LL, Claire TTR, Tchoumbou M and Helm B
Wood Warblers, an Afro-Palearctic migrant species, are declining steadily in Europe likely due to mortality outside their breeding grounds. However, little is known about their overwintering, and records about the sensitive life-cycle stage of moult in Africa are practically absent. To fill this gap, we report on moult of Wood Warblers captured over two winters (January-February) in 2019-2020 in Cameroon. We caught 14 individuals, of which 12 were monitored for flight feather moult. All inspected individuals showed advanced stages of flight feather renewal. Despite low sample sizes, Underhill-Zucchini moult models aptly explained variation in primary and secondary moult (  = 0.61). Estimated moult onset date was 26 December, completion date was 25 February, and moult duration was 61 days. These findings fit well with experimental data on the annual cycle and the timing of recently published migration tracks of Wood Warblers. Jointly, the data suggest that moult timing is set by an internal programme, which enables Wood Warblers to organise their multi-stage migration such that they reach suitable moulting habitat in time, and can depart in time with a fresh plumage for the breeding grounds. In our study, moult occurred during the peak of the dry season, which in Cameroon nonetheless shows high relative humidity. During our mist-netting on 28 cocoa plantations of varying shade cover, Wood Warblers were caught on 6 farms whose canopies were comparatively open. These data suggest that the birds encounter in Cameroon relatively stable climatic conditions for moult, and do not measurably prefer closed-canopy forests. Our findings are important, because successful moult increases survival prospects and because moult needs to be safely embedded in a migratory life cycle. Hence, information on moult timing and location is essential for identifying year-round vulnerabilities of Wood Warblers.
Excretion patterns of coccidian oocysts and nematode eggs during the reproductive season in Northern Bald Ibis ()
Frigerio D, Cibulski L, Ludwig SC, Campderrich I, Kotrschal K and Wascher CA
Individual reproductive success largely depends on the ability to optimize behaviour, immune function and the physiological stress response. We have investigated correlations between behaviour, faecal steroid metabolites, immune parameters, parasite excretion patterns and reproductive output in a critically endangered avian species, the Northern Bald Ibis (). In particular, we related haematocrit, heterophil/lymphocyte ratio, excreted immune-reactive corticosterone metabolites and social behaviour with parasite excretion and two individual fitness parameters, namely, number of eggs laid and number of fledglings. We found that the frequency of excretion of parasites' oocysts and eggs tended to increase with ambient temperature. Paired individuals excreted significantly more samples containing nematode eggs than unpaired ones. The excretion of nematode eggs was also significantly more frequent in females than in males. Individuals with a high proportion of droppings containing coccidian oocysts were more often preened by their partners than individuals with lower excretion rates. We observed that the more eggs an individual incubated and the fewer offspring fledged, the higher the rates of excreted samples containing coccidian oocysts. Our results confirm that social behaviour, physiology and parasite burden are linked in a complex and context-dependent manner. They also contribute background information supporting future conservation programmes dealing with this critically endangered species.
Common patterns in the molecular phylogeography of western palearctic birds: a comprehensive review
Pârâu LG and Wink M
A plethora of studies have offered crucial insights in the phylogeographic status of Western Palearctic bird species. However, an overview integrating all this information and analyzing the combined results is still missing. In this study, we compiled all published peer-reviewed and grey literature available on the phylogeography of Western Palearctic bird species. Our literature review indicates a total number of 198 studies, with the overwhelming majority published as journal articles ( = 186). In total, these literature items offer information on 145 bird species. 85 of these species are characterized by low genetic differentiation, 46 species indicate genetic variation but no geographic structuring i.e. panmixia, while 14 species show geographically distinct lineages and haplotypes. Majority of bird species inhabiting the Western Palearctic display genetic admixture. The glaciation cycles in the past few million years were pivotal factors in shaping this situation: during warm periods many species expanded their distribution range to the north over wide areas of Eurasia; whereas, during ice ages most areas were no longer suitable and species retreated to refugia, where lineages mixed.
From neurons to nests: nest-building behaviour as a model in behavioural and comparative neuroscience
Hall ZJ, Meddle SL and Healy SD
Despite centuries of observing the nest building of most extant bird species, we know surprisingly little about how birds build nests and, specifically, how the avian brain controls nest building. Here, we argue that nest building in birds may be a useful model behaviour in which to study how the brain controls behaviour. Specifically, we argue that nest building as a behavioural model provides a unique opportunity to study not only the mechanisms through which the brain controls behaviour within individuals of a single species but also how evolution may have shaped the brain to produce interspecific variation in nest-building behaviour. In this review, we outline the questions in both behavioural and comparative neuroscience that nest building could be used to address, summarize recent findings regarding the neurobiology of nest building in lab-reared zebra finches and across species building different nest structures, and suggest some future directions for the neurobiology of nest building.
Melanin-based plumage coloration and melanin content in organs in the barn owl
Roulin A, Dubey S, Ito S and Wakamatsu K
Although the evolutionary ecology of melanin pigments and melanin-based coloration has been studied in great details, particularly in birds, little is known about the function of melanin stored inside the body. In the barn owl , in which individuals vary in the degree of reddish pheomelanin-based coloration and in the size of black eumelanic feather spots, we measured the concentration in melanin pigments in seven organs. The eyes had by far the most melanin then the skin, pectoral muscle, heart, liver, trachea, and uropygial gland. The concentration in eumelanin was not necessarily correlated with the concentration in pheomelanin suggesting that their production can be regulated independently from each other. Redder barn owls had more pheomelanin in the skin and uropygial gland than white owls, while owls displaying larger black feather spots had more eumelanin in the skin than small-spotted owls. More data are required to evaluate whether melanin-based traits can evolve as an indirect response to selection exerted on melanin deposition in organs.