Cost, risk, and avoidance of inbreeding in a cooperatively breeding bird.

Inbreeding is often avoided in natural populations by passive processes such as sex-biased dispersal. But, in many social animals, opposite-sexed adult relatives are spatially clustered, generating a risk of incest and hence selection for active inbreeding avoidance. Here we show that, in long-tailed tits (Aegithalos caudatus), a cooperative breeder that risks inbreeding by living alongside opposite-sex relatives, inbreeding carries fitness costs and is avoided by active kin discrimination during mate choice. First, we identified a positive association between heterozygosity and fitness, indicating that inbreeding is costly. We then compared relatedness within breeding pairs to that expected under multiple mate-choice models, finding that pair relatedness is consistent with avoidance of first-order kin as partners. Finally, we show that the similarity of vocal cues offers a plausible mechanism for discrimination against first-order kin during mate choice. Long-tailed tits are known to discriminate between the calls of close kin and nonkin, and they favor first-order kin in cooperative contexts, so we conclude that long-tailed tits use the same kin discrimination rule to avoid inbreeding as they do to direct help toward kin.

Symmetry breaking in the embryonic skin triggers directional and sequential plumage patterning.

The development of an organism involves the formation of patterns from initially homogeneous surfaces in a reproducible manner. Simulations of various theoretical models recapitulate final states of natural patterns, yet drawing testable hypotheses from those often remains difficult. Consequently, little is known about pattern-forming events. Here, we surveyed plumage patterns and their emergence in Galliformes, ratites, passerines, and penguins, together representing the three major taxa of the avian phylogeny, and built a unified model that not only reproduces final patterns but also intrinsically generates shared and varying directionality, sequence, and duration of patterning. We used in vivo and ex vivo experiments to test its parameter-based predictions. We showed that directional and sequential pattern progression depends on a species-specific prepattern: an initial break in surface symmetry launches a travelling front of sharply defined, oriented domains with self-organising capacity. This front propagates through the timely transfer of increased cell density mediated by cell proliferation, which controls overall patterning duration. These results show that universal mechanisms combining prepatterning and self-organisation govern the timely emergence of the plumage pattern in birds.

Testing the developmental hypothesis of the HPA axis in a tropical passerine: Dampened corticosterone response and faster negative feedback in nestling lance-tailed manakins (Chiroxiphia lanceolata).

When vertebrates are exposed to stressors, the subsequent acute increase in glucocorticoids by the hypothalamic-pituitaryadrenal (HPA) axis triggers a suite of adaptive responses, including mobilization of stored energy and repression of non-essential processes. However, chronic exposure to high concentrations of glucocorticoids can lead to metabolic dysregulation, impaired immune function, and cognitive decline. In developing young, this hormonal stress response shows considerable variation. Generally, the physiological stress response of young of precocial species is comparable to that of adults, whereas offspring of altricial species exhibit an attenuated response compared to adults. The developmental hypothesis of the HPA axis proposes that the dampened stress response in dependent offspring is an adaptive response to avoid the negative effects of elevated glucocorticoids, particularly in altricial species where young lack the ability to mitigate stressful stimuli.We aimed to test the developmental hypothesis in a tropical avian species, the lance-tailed manakin (Chiroxiphia lanceolata). We predicted that nestlings of this altricial species should have a dampened corticosterone response, in both magnitude and duration, compared to that of adults. We also predicted that recently fledged hatch-year birds would display a response intermediate to that of adults and nestlings. We quantified circulating corticosterone levels in adults, recently fledged hatch-year birds, and 11-day-old nestlings using a standardized capture and restraint protocol. Nestlings showed a lower maximal corticosterone response and faster negative feedback compared to adults. Further, five post-fledging hatch-year birds showed a feedback response intermediate to those of nestlings and adults. However, we caution against generalizing about fledgling responses beyond this study due to the small sample (n = 5). Interestingly, lance-tailed manakin nestlings appear to return to baseline concentrations faster than nestlings of temperate species. These results support the developmental hypothesis of the HPA axis explaining variation in stress response. This study is the first to assess the development of the hormonal stress response in nestlings of a tropical bird, which is of interest because of our still-developing understanding of how tropical and temperate species differ physiologically. Finally, findings here underscore the importance of validating and adjusting sampling protocols that quantify nestling stress responses, as sampling timelines identified for adults may underestimate the magnitude of the nestling stress response.

Experimental Amelioration of Harsh Weather Speeds Growth and Development in a Tropical Montane Songbird.

AbstractOrganisms living at high elevations generally grow and develop more slowly than those at lower elevations. Slow montane ontogeny is thought to be an evolved adaptation to harsh environments that improves juvenile quality via physiological trade-offs. However, slower montane ontogeny may also reflect proximate influences of harsh weather on parental care and offspring development. We experimentally heated and protected nests from rain to ameliorate harsh montane weather conditions for mountain blackeyes (Chlorocharis emiliae), a montane songbird living at approximately 3,200 m asl in Malaysian Borneo. This experiment was designed to test whether cold and wet montane conditions contribute to parental care and postnatal growth and development rates at high elevations. We found that parents increased provisioning and reduced time spent warming offspring, which grew faster and departed the nest earlier compared with offspring from unmanipulated nests. Earlier departure reduces time-dependent predation risk, benefitting parents and offspring. These plastic responses highlight the importance of proximate weather contributions to broad patterns of montane ontogeny and parental care.

Tail colour signals performance in blue tit nestlings.

Indirect sexual selection arises when reproductive individuals choose their mates based on heritable ornaments that are genetically correlated to fitness. Evidence for genetic associations between ornamental colouration and fitness remains scarce. In this study, we investigate the quantitative genetic relationship between different aspects of tail structural colouration (brightness, hue and UV chroma) and performance (cell-mediated immunity, body mass and wing length) in blue tit (Cyanistes caeruleus) nestlings. In line with previous studies, we find low heritability for structural colouration and moderate heritability for performance measures. Multivariate animal models show positive genetic correlations between the three measures of performance, indicating quantitative genetic variation for overall performance, and tail brightness and UV chroma, two genetically independent colour measures, are genetically correlated with performance (positively and negatively, respectively). Our results suggest that mate choice based on independent aspects of tail colouration can have fitness payoffs in blue tits and provide support for the indirect benefits hypothesis. However, low heritability of tail structural colouration implies that indirect sexual selection on mate choice for this ornament will be a weak evolutionary force.

Elevated oxidative stress in pied flycatcher nestlings of eumelanic foster fathers under low rearing temperatures.

Striking variation in melanin coloration within natural populations is likely due to the different fitness outcomes of alternative phenotypes in varying environmental conditions. There are two types of melanin: eumelanins yield blackish hues, whereas pheomelanins yield reddish hues. The production of eumelanins requires low levels of glutathione (GSH), which is the most important intracellular antioxidant, whereas the production of pheomelanins requires high levels of GSH. We investigated the oxidative status of male pied flycatchers (Ficedula hypoleuca) with different degrees of melanin coloration under different temperatures during the nestling period. Moreover, we assessed the oxidative status of offspring in relation to their biological or foster father's melanin coloration and ambient temperature. To separate offspring genotype effects and paternal effects in different temperatures, we used a partial cross-foster design. The temperature differently affected the oxidative status of differently colored male pied flycatchers and their foster offspring. When the weather was relatively cold, black males had higher glutathione S-transferase levels compared with brown males, indicating enhanced stress in black males. Foster offspring of black males had a lower ratio between reduced and oxidized GSH followed by higher total amount of GSH than foster offspring of brown males. Thus, foster offspring of black males seem to suffer from oxidative stress under relatively cold weather compared with those of brown males, and vice versa under relatively warm weather. Although differently colored males experienced changes in their oxidative status under different temperatures, the link between paternal melanin coloration and offspring oxidative stress appears to be environmentally induced.

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird.

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long-term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg-laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year-to-year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival-breeding interval (earlier breeding). Finally, mark-recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.

Do parasites and antioxidant availability affect begging behaviour, growth rate and resistance to oxidative stress?

Early-life trade-offs faced by developing offspring can have long-term consequences for their future fitness. Young offspring use begging displays to solicit resources from their parents and have been selected to grow fast to maximize survival. However, growth and begging behaviour are generally traded off against self-maintenance. Oxidative stress, a physiological mediator of life-history trade-offs, may play a major role in this trade-off by constraining, or being costly to, growth and begging behaviour. Yet, despite implications for the evolution of life-history strategies and parent-offspring conflicts, the interplay between growth, begging behaviour and resistance to oxidative stress remains to be investigated. We experimentally challenged wild great tit (Parus major) offspring by infesting nests with a common ectoparasite, the hen flea (Ceratophyllus gallinae), and simultaneously tested for compensating effects of increased vitamin E availability, a common dietary antioxidant. We further quantified the experimental treatment effects on offspring growth, begging intensity and oxidative stress. Flea-infested nestlings of both sexes showed reduced body mass during the first half of the nestling phase, but this effect vanished short before fledging. Begging intensity and oxidative stress of both sexes were unaffected by both experimental treatments. Feeding rates were not affected by the experimental treatments, but parents of flea-infested nests fed nestlings with a higher proportion of caterpillars, the main source of antioxidants. Additionally, female nestlings begged significantly less than males in control nests, whereas both sexes begged at similar rates in vitamin E-supplemented nests. Our study shows that a parasite exposure does not necessarily affect oxidative stress levels or begging intensity, but suggests that parents can compensate for negative effects of parasitism by modifying food composition. Furthermore, our results indicate that the begging capacity of the less competitive sex is constrained by antioxidant availability.

Oviposition drives hatching order and developmental disparities with brood mates.

Birth or hatching order can affect fitness. It has long been assumed that the order in which a bird's egg is laid (its oviposition) and first exposed to incubation relative to other eggs in the nest determines the order in which it hatches and the subsequent effects on development and survival. To my knowledge, this cause of hatching order has not been tested while controlling for laying-order effects on egg composition. Like those of many species, the last-laid eggs in clutches of the Lincoln's sparrow ( Melospiza lincolnii) hatch last and produce nestlings with reduced growth and survival rates. Last-hatched Lincoln's sparrows also develop wider bills that are associated with reduced attractiveness of adult courtship songs. Using wild, free-ranging Lincoln's sparrows, I replaced the first or fourth freshly laid egg in four-egg host nests with a freshly laid first egg from a donor nest. I discovered that the experimental elevation of oviposition-which controlled for egg composition-delayed hatching, inhibited growth and widened bill gape, thus confirming the mechanism for hatching order and revealing a potential role for oviposition in reproductive success. Similar effects of postnatal family position could affect fitness in other taxa, as well.

Effects of Experimental Lead Exposure on Testis of the Chestnut Capped Blackbird Chrysomus ruficapillus.

Lead (Pb) effects on testis histology, as well as sperm quality and oxidative status were evaluated in male Chestnut Capped Blackbird (Chrysomus ruficapillus). Wild blackbirds were captured, immediately sampled (field group) or kept in captivity and treated with a single intraperitoneal injection of saline solution (control) or saline solution with Pb acetate (50 or 100 mg/kg Pb). Seven days after injection, whole blood, ductus deferens and testis samples were collected. Increased Pb concentrations were observed in whole blood and testis of Pb-exposed blackbirds with respect to those from field and control blackbirds. Sperm cells of Pb-exposed blackbirds showed loss of membrane integrity, mitochondrial functionality, and DNA integrity. Also, oxidative damage was observed in testis of blackbirds injected with 100 mg/kg Pb. These findings indicate that Pb is accumulated in testis of C. ruficapillus, inducing severe morphological and biochemical injury that can compromise the reproductive performance of male blackbirds. Although the exposure scenario (Pb acetate, high dosage and intraperitoneal injection) tested in the present study would likely not occur in the wild, it was adequate to show potential and relevant toxic effects of Pb in wild birds.

Effects of experimentally manipulated yolk thyroid hormone levels on offspring development in a wild bird species.

Maternal effects are a crucial mechanism in a wide array of taxa to generate phenotypic variation, thereby affecting offspring development and fitness. Maternally derived thyroid hormones (THs) are known to be essential for offspring development in mammalian and fish models, but have been largely neglected in avian studies, especially in respect to natural variation and an ecological context. We studied, for the first time in a wild species and population, the effects of maternally derived THs on offspring development, behavior, physiology and fitness-related traits by experimental elevation of thyroxine and triiodothyronine in ovo within the physiological range in great tits (Parus major). We found that elevated yolk TH levels had a sex-specific effect on growth, increasing male and decreasing female growth, relative to controls, and this effect was similar throughout the nestling period. Hatching or fledging success, motor coordination behavior, stress reactivity and resting metabolic rate were not affected by the TH treatment. We conclude that natural variation in maternally derived THs may affect some offspring traits in a wild species. As this is the first study on yolk thyroid hormones in a wild species and population, more such studies are needed to investigate its effects on pre-hatching development, and juvenile and adult fitness before generalizations on the importance of maternally derived yolk thyroid hormones can be made. However, this opens a new, interesting avenue for further research in the field of hormone mediated maternal effects.

Brood size constrains the development of endothermy in blue tits.

Altricial birds are unable to maintain body temperature when exposed to low ambient temperatures during the first days after hatching. Thermoregulatory capacity begins to form as postnatal development progresses, and eventually nestlings become homeothermic. Several factors may influence this development at both the level of the individual and the level of the whole brood, but to our knowledge no studies have focused on the effect of brood size per se on the development of endothermy in individual nestlings. We performed cooling experiments on blue tit (Cyanistes caeruleus) nestlings in the field, to study how different experimental brood sizes affected the development of endothermy in individual nestlings and the thermal environment experienced by the whole brood in the nest. Nestlings from all experimental brood sizes showed a decrease in cooling rate as they grew older, but birds from reduced broods showed an earlier onset of endothermy compared with nestlings from enlarged and control broods. This difference manifested during early development and gradually disappeared as nestlings grew older. The thermal environment in the nests differed between treatments during nestling development, such that nest temperature in reduced broods was lower than that in enlarged broods during all days and during nights at the end of the experimental period. We suggest that the development of endothermy in blue tit nestlings is not ontogenetically fixed, but instead may vary according to differences in developmental, nutritional and thermal conditions as determined by brood size.

Dietary flavonoids advance timing of moult but do not affect redox status of juvenile blackbirds (Turdus merula).

Flavonoids are the most abundant plant polyphenols, widely occurring in fruits and berries, and show a strong antioxidant activity in vitro Studies of avian species feeding on berries suggest that dietary flavonoids have health-promoting effects and may enhance the expression of melanin-based plumage traits. These effects are probably mediated by the antioxidant activity of flavonoids. However, the effect of dietary flavonoids on oxidative status has never been investigated in any bird species. We analysed the effects of dietary flavonoids on blood non-enzymatic antioxidants and protein oxidative damage of juvenile European blackbirds (Turdus merula). In addition, we analysed the effects of flavonoid-enriched diet on body condition and on the timing of moult from juvenile to adult plumage. Dietary flavonoids did not significantly affect redox status but significantly advanced the onset of moult, hastening plumage development. Moulting birds showed higher protein oxidative damage compared with those that had not yet started moulting. The probability of initiating moult after 40 days of dietary treatment was higher for birds with low circulating levels of oxidizing agents and high glutathione concentration. The metabolization of flavonoids could have altered their redox potential, resulting in no net effects on redox status. However, flavonoid consumption before and during moult may contribute to enhance plumage development. Moreover, our findings suggest that moulting feathers may result in redox imbalance. Given their effect on moult and growth of melanin-rich feathers, fruit flavonoids may have contributed to shape plant fruiting time in relation to fruit consumption preferences by birds.

Long-term consequences of high incubation temperature in a wild bird population.

Because incubation by birds is energetically costly, parents frequently trade off investment in incubation against self-maintenance. This can be manifested by a reduction in incubation temperature, which comes at high somatic costs for nestlings. The extent to which these costs constrain fitness is poorly understood. We incubated wild blue tit clutches at three biologically relevant temperatures and subsequently recorded winter survival and survival to the breeding season. Fledglings from the coldest treatment (35.0°C) survived less well than other fledglings, but the proportion of winter and breeding survivors did not differ significantly between treatments. However, survival probability in both seasons increased with body mass at fledging in birds from low and mid incubation temperatures, but decreased with fledging body mass in the high-temperature treatment. Mid-temperature nestlings were heavier as adults, weighing 7% more than low- and high-temperature survivors. Thus, high incubation temperature can be beneficial in the short term, but costs of accelerated embryonic development may equal those of protracted development in the long term. Such hidden consequences of faster development could maintain natural selection for average incubation temperature.

Differential prevalence and diversity of haemosporidian parasites in two sympatric closely related non-migratory passerines.

Haemosporidian parasites infecting birds show distinct heterogeneity in their distribution among host species. However, despite numerous studies on the prevalence and diversity of parasite communities across species, very little is known on patterns of differences between them. Such data is lacking because up to date the majority of studies explored the patterns of variation in infections in different years, different time of sampling within a year or a breeding cycle, different study sites or was based on a small sample size, all of which may affect the estimates of prevalence and parasite diversity. Here, the prevalence, richness and diversity of haemosporidian parasites from the genera Plasmodium and Haemoproteus were studied in two closely related non-migratory hole-nesting passerines: Great Tits and Blue Tits. Birds were sampled in sympatrically breeding populations during two seasons at the same stage of their breeding cycle - late nestling care. Great Tits were more prevalently infected with Plasmodium and Haemoproteus parasites (97·1 vs 71·2%), harboured a higher proportion of multiple infections (26·2 vs 3·2%) and had a more diverse parasite community (11 vs 5 parasite lineages) than Blue Tits. Observed differences between two host species are discussed with reference to their breeding densities and immunological and behavioural characteristics.

Local weather conditions have complex effects on the growth of blue tit nestlings.

Adverse weather conditions are expected to result in impaired nestling development in birds, but empirical studies have provided equivocal support for such a relationship. This may be because the negative effects of adverse weather conditions are masked by parental effects. Globally, ambient temperatures, rainfall levels and wind speeds are all expected to increase in a changing climate and so there is a need for a better understanding of the relationship between weather conditions and nestling growth. Here, we describe a correlative study that examined the relationships between local temperatures, rainfall levels and wind speeds and the growth of individual blue tit (Cyanistes caeruleus) nestlings in relation to their hatching order and sex. We found that changes in a range of morphological characters were negatively related to both temperature and wind speed, but positively related to rainfall. These patterns were further influenced by the hatching order of the nestlings but not by nestling sex. This suggests that the predicted changes in local weather conditions may have complex effects on nestling growth, but that parents may be able to mitigate the adverse effects via adaptive parental effects. We therefore conclude that local weather conditions have complex effects on avian growth and the implications for patterns of avian growth in a changing climate are discussed.

Weathering the storm: parental effort and experimental manipulation of stress hormones predict brood survival.

BACKGROUND: Unpredictable and inclement weather is increasing in strength and frequency, challenging organisms to respond adaptively. One way in which animals respond to environmental challenges is through the secretion of glucocorticoid stress hormones. These hormones mobilize energy stores and suppress non-essential physiological and behavioral processes until the challenge passes. To investigate the effects of glucocorticoids on reproductive decisions, we experimentally increased corticosterone levels (the primary glucocorticoid in birds) in free-living female tree swallows, Tachycineta bicolor, during the chick-rearing stage. Due to an unprecedented cold and wet breeding season, 90 % of the nests in our study population failed, which created a unique opportunity to test how challenging environmental conditions interact with the physiological mechanisms underlying life-history trade-offs. RESULTS: We found that exogenous corticosterone influenced the regulation of parental decisions in a context-dependent manner. Control and corticosterone-treated females had similar brood failure rates under unfavorable conditions (cold and rainy weather), but corticosterone treatment hastened brood mortality under more favorable conditions. Higher female nest provisioning rates prior to implantation were associated with increased probability of brood survival for treatment and control groups. However, higher pre-treatment male provisioning rates were associated with increased survival probability in the control group, but not the corticosterone-treated group. CONCLUSIONS: These findings reveal complex interactions between weather, female physiological state, and partner parental investment. Our results also demonstrate a causal relationship between corticosterone concentrations and individual reproductive behaviors, and point to a mechanism for why naturally disturbed populations, which experience multiple stressors, could be more susceptible and unable to respond adaptively to changing environmental conditions.

Conflict between biotic and climatic selective pressures acting on an extended phenotype in a subarctic, but not temperate, environment.

Climatic selective pressures are thought to dominate biotic selective pressures at higher latitudes. However, few studies have experimentally tested how these selective pressures differentially act on traits across latitudes because traits can rarely be manipulated independently of the organism in nature. We overcame this challenge by using an extended phenotype-active bird nests-and conducted reciprocal transplant experiments between a subarctic and temperate site, separated by 14° of latitude. At the subarctic site, biotic selective pressures (nest predation) favoured smaller, non-local temperate nests, whereas climatic selective pressures (temperature) favoured larger local nests, particularly at colder temperatures. By contrast, at the temperate site, climatic and biotic selective pressures acted similarly on temperate and subarctic nests. Our results illustrate a functional trade-off in the subarctic between nest morphologies favoured by biotic versus climatic selective pressures, with climate favouring local nest morphologies. At our temperate site, however, allocative trade-offs in the time and effort devoted to nest construction favour smaller, local nests. Our findings illustrate a conflict between biotic and climatic selective pressures at the northern extremes of a species geographical range, and suggest that trade-offs between trait function and trait elaboration act differentially across latitude to create broad geographic variation in traits.

Is oxidative status influenced by dietary carotenoid and physical activity after moult in the great tit (Parus major)?

In the context of sexual and natural selection, an allocation trade-off for carotenoid pigments may exist because of their obligate dietary origin and their role both in the antioxidant and immune systems and in the production of coloured signals in various taxa, particularly birds. When birds have expended large amounts of carotenoids to feather growth such as after autumn moult, bird health and oxidative status might be more constrained. We tested this hypothesis in a bird species with carotenoid-based plumage colour, by manipulating dietary carotenoids and physical activity, which can decrease antioxidant capacity and increase reactive oxygen metabolite (ROM) concentration. Great tits were captured after moult and kept in aviaries, under three treatments: physical handicap and dietary supplementation with carotenoids, physical handicap and control diet, and no handicap and control diet. We measured plasma composition (antioxidant capacity, ROM concentration, and vitamin A, vitamin E and total carotenoid concentrations), immune system activation (blood sedimentation) and stress response (heterophil/lymphocyte ratio) and predicted that handicap treatment should influence these negatively and carotenoid supplementation positively. Coloration of yellow feathers was also measured. Carotenoid supplementation increased total plasma carotenoid concentration, decreased feather carotenoid chroma and marginally increased ROM concentration. Handicap increased blood sedimentation only in males but had no clear influence on oxidative stress, which contradicted previous studies. Further studies are needed to investigate how physical activity and carotenoid availability might interact and influence oxidative stress outside the moult period, and their combined potential influence on attractiveness and reproductive investment later during the breeding season.

Adult and hatch-year blackpoll warblers exhibit radically different regional-scale movements during post-fledging dispersal.

Using a broad-scale automated telemetry array, we explored post-fledging movements of blackpoll warblers breeding in Atlantic Canada. We sought to determine the full spatial scale of post-fledging dispersal, to assess support for three hypotheses for regional-scale post-fledging movement, and to determine whether learning influenced movement during this period. We demonstrated that both young and adults moved over distances more than 200 km prior to initiating migration. Adults moved southwest, crossing the Gulf of Maine (GOM), consistent with the commencement of migration hypothesis. Hatch-year birds exhibited less directional movements constrained geographically by the GOM. Their movements were most consistent with exploration hypotheses--that young birds develop a regional-scale map to aid in habitat selection, natal dispersal and subsequent migrations.