Post-breeding Migration Research Articles

Cue-conflict experiments between magnetic and visual cues in dunlin Calidris alpina and curlew sandpiper Calidris ferruginea

Despite our extensive knowledge on various aspects of their lives, there has been limited investigation into the hierarchical relationships among different compass systems in shorebirds. The aim of this study was to investigate the relationship between magnetic and celestial compasses in two species of shorebirds, the curlew sandpiper (Calidris ferruginea; pre-breeding migration) and the dunlin (Calidris alpina; post-breeding migration) using cue-conflict experiments. Birds were captured in a Mediterranean stopover site, after which their magnetic orientation was determined under simulated overcast conditions at sunset using modified Emlen funnels fitted with infrared video cameras. Birds that demonstrated a well-defined directional preference were then exposed over two sunsets to conflicting directional information between the local geomagnetic field and the ±90° shifted band of maximum polarisation. These individuals were tested again for magnetic orientation at sunset in the same conditions as previous test, to determine whether their directional choices had changed after the cue-conflict. Our results showed that individuals from both species did not recalibrate their magnetic compass from visual cues after the cue-conflict, even though at least dunlins did not appear to completely disregard the information derived from celestial cues. This study is one of the few experimental studies on the migratory orientation of Charadriiformes and on the hierarchical relationships between the different compasses used by these birds during their extensive migratory movements. Migrating birds are able to use different compass mechanisms based on geomagnetic or celestial cues, and it seems reasonable to hypothesise that birds calibrate their various compasses to maintain the correct direction especially when the directional information does not agree. The hierarchy among different compasses has been studied largely on night migrating passerines, but it is still poorly understood. We investigated the hierarchy among geomagnetic and celestial cues (band of maximum polarisation) in two species of Charadriiformes by means of cue-conflict experiments. Our result showed that the geomagnetic cues have a dominant role in the orientation mechanisms of the studied species, even though the information derived from celestial cues did not appear to be completely disregarded.

Abundance, occurrence, and seasonality of the Subtropical Doradito (Pseudocolopteryx acutipennis) on the coast of Brazil

ABSTRACT The Subtropical Doradito (Pseudocolopteryx acutipennis) is a small tyrannid that breeds in shrubby marshlands of South America. Post-breeding migration occurs in both mountain and lowland populations, but little is known of the destination(s) of these birds. Subtropical Doraditos have recently been found during the southern winter, or non-breeding season, in several localities in southern Amazonian Brazil and the Pantanal, with four reports from coastal Brazil. Our observations and censuses in coastal marshes of southeast and southern Brazil have produced eight records of Subtropical Doradito in the states of Parana and Sao Paulo. In light of comparison with other Brazilian records compiled for this study, we suggest that coastal winterers are likely latitudinal migrants from geographically restricted breeding grounds in the Pampas of Argentina. In addition, we discuss conservation problems for the Subtropical Doradito in coastal wintering areas.

Migration flyway of the Mediterranean breeding Lesser Crested Tern Thalasseus bengalensis emigratus

The Lesser Crested Tern Thalasseus bengalensis emigratus breeding population in the Mediterranean is found exclusively in Libya, on the two coastal islands of Gara and Elba and one wetland on the mainland coast at Benghazi. In order to improve knowledge of the species migration to wintering quarters in West Africa, a ringing programme was conducted from 2006–2008 and 2009–2012. From a total of 1 354 nestlings ringed using metal and/or colour rings, 64 were recovered along their flyway and in their wintering range, representing 6.9% of birds ringed with both colour and metal rings. This provided the opportunity to collect information on post-natal movements (staging and wintering ranges), breeding philopatry and recruitment, in addition to a preliminary estimate of their migration journey duration. This paper indicates sighting and recovery distributions in space and time, highlighting the important areas for the species during its journey between breeding and wintering sites. The findings indicate that several areas where ringed terns stop-over during pre- and post-breeding migration journeys are not protected, causing an additional threat to their survival, as some wintering areas are also not protected. Conservation of this highly localised and threatened population needs not only to address protection at breeding sites but also at migratory stop-overs and wintering strongholds.

Post-breeding migration routes of marine turtles from Bonaire and Klein Bonaire, Caribbean Netherlands

The management of small rookeries is key to conserving the regional genetic diversity of marine turtle populations and requires knowledge on population connectivity between breeding and foraging areas. To elucidate the geographic scope of the populations of marine turtles breeding at Bonaire and Klein Bonaire (Caribbean Netherlands) we examined the post-breeding migratory behavior of 5 female loggerheads Caretta caretta, 4 female green turtles Chelonia mydas, and 2 male and 13 female hawksbill turtles Eretmochelys imbricata during the years 2004-2013. After leaving Bonaire, the 24 tracked turtles frequented foraging grounds in 10 countries. The distances swum from Bonaire to the foraging areas ranged from 608 to 1766 km for loggerhead turtles, 198 to 3135 km for green turtles, and 197 to 3135 km for hawksbill turtles, together crossing the waters of 19 countries. Males represented the minority in this study, but we made 2 key observations that require further research: males remained in the vicinity of the breeding area for 3-5 mo, which is 2-5 times longer than females, and males migrated greater distances than previously recorded. Although the turtles dispersed widely across the Caribbean, there appeared to be 2 benthic foraging areas of particular importance to all 3 species of marine turtles breeding at Bonaire, namely the shallow banks east of Nicaragua and Honduras (n = 8 tracked turtles) and Los Roques, Venezuela (n = 3). Marine turtles breeding at Bonaire face threats from legal turtle harvesting, illegal take, and bycatch in the waters that they traverse across the Caribbean.

Use of artificial intertidal sites by birds in a Mediterranean lagoon and their importance for wintering and migrating waders

Man-made habitats provide suitable nesting, resting and feeding habitats for many birds at coastal sites. Despite intensive study outside Europe, very few data are available to date on the bird communities that exploit artificial intertidal sites along the European coasts. Between July 2009 and December 2010, 32 ornithological surveys were performed at six artificial intertidal sites in the lagoon of Venice (Italy). Overall, 101 species (with 23,399 birds) were observed, about a third of those occurring in the lagoon of Venice; 58 of these species were of conservation concern. Ten species comprised about 80 % of the total; bird abundance was the highest during the post-breeding migration and wintering period. Multivariate analysis identified similarity among sites, seasons and observed behaviours. Dredge islands were mostly used by waders as foraging sites, when surrounding tidal flats were still flooded. In winter the six dredged islands supported about 2 % of the dunlins Calidris alpina occurring in the whole lagoon of Venice and about 10 % of the grey plovers Pluvialis squatarola. Since the study sites comprised just the 6 % of the total dredge island area available in 2009–2010, the results indicate as large fractions of these shorebird populations might exploit the dredge island habitats for foraging. Among the newly created habitats, intertidal ponds and inner tidal flats were the most heavily used by birds, followed by dykes and mounds with ruderal vegetation. Maintenance of the habitat mosaic is a need for a conservation- based management of the dredge islands.

Post-breeding migration of Night Herons Nycticorax nycticorax tracked by GPS/GSM transmitters

AbstractThe migratory strategy of herons is poorly understood: only that of Purple Heron has been well studied. The migratory movements of three Night Herons from the Upper Vistula Valley (Poland) were studied using a GPS/GSM-tracking system in 2012. The results document a previously unknown autumn migration strategy of Night Herons with long stopovers en route, unlike that of Purple Herons. The different migration strategies of Purple Herons and Night Herons could be due to differences in wing-loadings and foraging ecology.

Annual movement patterns of endangered ivory gulls: the importance of sea ice.

The ivory gull (Pagophila eburnea) is an endangered seabird that spends its entire year in the Arctic environment. In the past three decades, threats from various sources have contributed to a >70% decline in Canada. To assess the annual habitat needs of this species, we attached satellite transmitters to 12 ivory gulls on Seymour Island, Nunavut in 2010, which provided up to four breeding seasons of tracking data. Analysis of migratory behaviour revealed considerable individual variation of post-breeding migratory route selection. Ivory gulls traveled a median of 74 days during post-breeding migration, but only 18 days during pre-breeding migration. In contrast to predictions, ivory gulls did not use the Greenland coast during migratory periods. Ivory gulls overwintered near the ice edge in Davis Strait, but also used the Labrador Sea in late February and March. We suggest that the timing of formation and recession and extent of sea ice plays a large role in ivory gull distribution and migratory timing.

New population estimates of a critically endangered species, the Balearic Shearwater Puffinus mauretanicus, based on coastal migration counts

SummaryThe Balearic Shearwater Puffinus mauretanicus is considered one of the most threatened seabirds in the world, with the breeding population thought to be in the range of 2,000–3,200 breeding pairs, from which global population has been inferred as 10,000 to 15,000 birds. To test whether the actual population of Balearic Shearwaters is larger than presently thought, we analysed the data from four land-based census campaigns of Balearic Shearwater post-breeding migration through the Strait of Gibraltar (mid-May to mid-July 2007–2010). The raw results of the counts, covering from 37% to 67% of the daylight time throughout the migratory period, all revealed figures in excess of 12,000 birds, and went up to almost 18,000 in two years. Generalised Additive Models were used to estimate the numbers of birds passing during the time periods in which counts were not undertaken (count gaps), and their associated error. The addition of both counted and estimated birds reveals figures of between 23,780 and 26,535 Balearic Shearwaters migrating along the north coast of the Strait of Gibraltar in each of the four years of our study. The effects of several sources of bias suggest a slight potential underestimation in our results. These figures reveal the urgent need to reformulate the population viability analysis for the species, and then if necessary reconsider its conservation status.

Ecology of Aquatic WarblersAcrocephalus paludicolain a Fall Stopover Area on the Atlantic Coast of France

The Aquatic Warbler Acrocephalus paludicola is the only endangered songbird in continental Europe. This trans-Saharan migratory bird significantly transits along the French Atlantic coastline during post-breeding migration and the right bank of the Gironde estuary has been identified as an important stopover site. We studied the spatial occupancy strategies of stationing individuals during August of three successive years (2010–2012). We characterized habitat use by radio-tracking individuals revealing relatively small foraging ranges (6.6 ± 2.6 ha on average) with only 1 ha actually exploited (core area), and a relatively high habitat fragmentation rate. Capture-mark-recapture analyses assessed the average stopover duration of individuals and body-mass variations during their stopover. The estimated average stopover duration was 6.46 ± 0.46 days (95% confidence interval: 4.4–9.6). Lean birds tended to forage significantly more than stout birds: on average, they gain 2.81 ± 0.89% of their initial mass each day whereas stout birds only gain 0.12 ± 0.56%. Analyses of droppings characterized the local diet. We noticed that Aquatic Warbler preferentially used partially-flooded or flooded habitats with heterogeneous and rather low vegetation, such as bulrush beds or bulrush beds mixed with reed beds. Orthoptera, Araneae and Hymenoptera represented the largest contributions to the consumed biomass (64.7%, 13.4% and 8.9% respectively). The importance of the fuel deposition rate of lean birds reflects the importance of the estuary as a stopover site for the species. It means that the available resources allow birds to replenish and continue their migration route. However, the sustainability of the site's functionality is questioned because of the evolution of habitats (erosion, rise in water levels and changes in food web).

Why water birds forage at night: a test using black‐tailed godwits Limosa limosa during migratory periods

Many migratory water birds are known to feed both during day and night outside the breeding season, but the underlying factors and mechanisms determining this foraging pattern are poorly understood. We addressed this topic by comparing both diurnal and nocturnal foraging activity (FA) and metabolizable energy intake rate (MEIR) in migrating black‐tailed godwits Limosa limosa staging in two different habitats, rice fields and coastal salt pans. Black‐tailed godwits staging in rice fields during pre‐breeding migration fed on rice seeds, and only foraged during the daylight period (FA: 81.89 ± 3.03%; MEIR: 1.15 ± 0.03 kJ · min−1). Daily energy consumption (DEC) of godwits relying on seeds was enough to meet the theoretical daily energy expenditure (DEE). In contrast, black‐tailed godwits staging in salt pans during post‐breeding migration fed on chironomid larvae, and they foraged during both daylight (FA: 67.36 ± 4.30%; MEIR: 0.27 ± 0.01 kJ · min−1) and darkness (FA: 69.89 ± 6.89%; MEIR: 0.26 ± 0.00 kJ · min−1). Nocturnal energy intake contributed 31.7% to DEC, the latter being insufficient to fully meet DEE. Our findings give empirical support to the view that diurnal foraging is the norm in many migratory water birds outside the breeding season, and nocturnal foraging occurs when the daily energy requirements are not met during the daylight period, supporting the supplementary food hypothesis.

A Space Oddity: Geographic and Specific Modulation of Migration in Eudyptes Penguins

Post-breeding migration in land-based marine animals is thought to offset seasonal deterioration in foraging or other important environmental conditions at the breeding site. However the inter-breeding distribution of such animals may reflect not only their optimal habitat, but more subtle influences on an individual’s migration path, including such factors as the intrinsic influence of each locality’s paleoenvironment, thereby influencing animals’ wintering distribution. In this study we investigated the influence of the regional marine environment on the migration patterns of a poorly known, but important seabird group. We studied the inter-breeding migration patterns in three species of Eudyptes penguins (E. chrysolophus, E. filholi and E. moseleyi), the main marine prey consumers amongst the World’s seabirds. Using ultra-miniaturized logging devices (light-based geolocators) and satellite tags, we tracked 87 migrating individuals originating from 4 sites in the southern Indian Ocean (Marion, Crozet, Kerguelen and Amsterdam Islands) and modelled their wintering habitat using the MADIFA niche modelling technique. For each site, sympatric species followed a similar compass bearing during migration with consistent species-specific latitudinal shifts. Within each species, individuals breeding on different islands showed contrasting migration patterns but similar winter habitat preferences driven by sea-surface temperatures. Our results show that inter-breeding migration patterns in sibling penguin species depend primarily on the site of origin and secondly on the species. Such site-specific migration bearings, together with similar wintering habitat used by parapatrics, support the hypothesis that migration behaviour is affected by the intrinsic characteristics of each site. The paleo-oceanographic conditions (primarily, sea-surface temperatures) when the populations first colonized each of these sites may have been an important determinant of subsequent migration patterns. Based on previous chronological schemes of taxonomic radiation and geographical expansion of the genus Eudyptes, we propose a simple scenario to depict the chronological onset of contrasting migration patterns within this penguin group.

Pandemic H1N1 influenza isolated from free-ranging Northern Elephant Seals in 2010 off the central California coast.

Interspecies transmission of influenza A is an important factor in the evolution and ecology of influenza viruses. Marine mammals are in contact with a number of influenza reservoirs, including aquatic birds and humans, and this may facilitate transmission among avian and mammalian hosts. Virus isolation, whole genome sequencing, and hemagluttination inhibition assay confirmed that exposure to pandemic H1N1 influenza virus occurred among free-ranging Northern Elephant Seals (Mirounga angustirostris) in 2010. Nasal swabs were collected from 42 adult female seals in April 2010, just after the animals had returned to the central California coast from their short post-breeding migration in the northeast Pacific. Swabs from two seals tested positive by RT-PCR for the matrix gene, and virus was isolated from each by inoculation into embryonic chicken eggs. Whole genome sequencing revealed greater than 99% homology with A/California/04/2009 (H1N1) that emerged in humans from swine in 2009. Analysis of more than 300 serum samples showed that samples collected early in 2010 (n = 100) were negative and by April animals began to test positive for antibodies against the pH1N1 virus (HI titer of ≥1∶40), supporting the molecular findings. In vitro characterizations studies revealed that viral replication was indistinguishable from that of reference strains of pH1N1 in canine kidney cells, but replication was inefficient in human epithelial respiratory cells, indicating these isolates may be elephant seal adapted viruses. Thus findings confirmed that exposure to pandemic H1N1 that was circulating in people in 2009 occurred among free-ranging Northern Elephant Seals in 2010 off the central California coast. This is the first report of pH1N1 (A/Elephant seal/California/1/2010) in any marine mammal and provides evidence for cross species transmission of influenza viruses in free-ranging wildlife and movement of influenza viruses between humans and wildlife.

Migration Routes and Staging Areas of Trans-Saharan Turtle Doves Appraised from Light-Level Geolocators

The identification of migration routes, wintering grounds and stopover sites are crucial issues for the understanding of the Palearctic-African bird migration system as well as for the development of relevant conservation strategies for trans-Saharan migrants. Using miniaturized light-level geolocators we report a comprehensive and detailed year round track of a granivorous trans-Saharan migrant, the European Turtle Dove (Streptopelia turtur). From five recovered loggers, our data provide new insights on migratory journeys and winter destinations of Turtle Doves originating from a breeding population in Western France. Data confirm that Turtle Doves wintered in West Africa. The main wintering area encompassed Western Mali, the Inner Delta Niger and the Malian/Mauritanian border. Some individuals also extended their wintering ranges over North Guinea, North-West of Burkina Faso and the Ivory-Coast. Our results reveal that all individuals did not spend the winter period at a single location; some of them experienced a clear eastward shift of several hundred kilometres. We also found evidence for a loop migration pattern, with a post-breeding migration flyway lying west of the spring route. Finally, we found that on their way back to breeding grounds Turtle Doves needed to refuel after crossing the Sahara desert. Contrary to previous suggestions, our data reveal that birds used stopover sites for several weeks, presumably in Morocco and North Algeria. This later finding is a crucial issue for future conservation strategies because environmental conditions on these staging areas might play a pivotal role in population dynamics of this declining species.

Piecing together the epic transoceanic migration of the Long-tailed Cuckoo (Eudynamys taitensis): an analysis of museum and sighting records

Long-tailed Cuckoos (Eudynamys taitensis) breed in New Zealand and winter in an arc of small Pacific islands extending 11 000 km from west to east. To understand this migration better, we compiled records of Cuckoos in time (month) and space (latitude, longitude), using museum voucher specimens and literature records. At the start of the breeding season (October-December) virtually all birds in New Zealand are in adult plumage; non-adult Cuckoos are almost entirely absent until late summer and autumn. In the northward post-breeding migration, adults move first and non-adults show a lag of up to 3 months in the timing of their departure. Non-adults are scarce after June in the Pacific wintering islands, and transitionals (with plumage between that of adults and non-adults) are very scarce after September, suggesting that they quickly moult into adult plumage. From June to October the average longitude of Cuckoos wintering north of New Zealand decreases progressively, consistent with a movement westward. This suggests that a portion of the Cuckoo population may undertake an anticlockwise loop-migration. This may also partly explain the unusual distribution of the species fanning out from the breeding to the wintering grounds.

Post-breeding migration of the Swallow, Hirundo rustica, along the coast of Lazio

We have studied (1995-2006) the autumn migration of the Swallow along the coast of Lazio. The individuals migrate preferably towards SE especially when the winds blow from S and SW at a speed between 10 and 12 km/h. Since 2003 the passages occur about ten days earlier.

Different Means to the Same End: Long-Distance Migrant Seabirds from Two Colonies Differ in Behaviour, Despite Common Wintering Grounds

Although seabirds that are trans-equatorial migrants show apparently broad overlap among populations in the non-breeding season, such large-scale pattern may conceal subtle but nevertheless key differences in migratory behaviour. These specializations could reflect adaptation to different environments during the breeding season, carry-over effects from the breeding to the nonbreeding period, or asymmetries in competitive ability of birds of different origin. We compared the migratory and wintering behaviour of Cory's shearwaters Calonectris diomedea nesting in Berlengas and in the Selvagens, two colonies in contrasting oceanographic environments, separated by ca. 1200 km. Although no differences were found in winter distribution, there was a marked divergence in timing, route and the use of staging areas during the postbreeding (autumn) migration. Birds from Berlengas typically travelled to oceanic waters in the North Atlantic for an extended stopover, whereas those from Selvagens rarely did so. In the South Atlantic, birds from Selvagens spent more time in flight, perhaps because they had higher energy and nutrient requirements for feather replacement compared to birds from Berlengas, which moult more flight feathers during breeding. Stable isotope analyses of feathers suggested that this variation in activity patterns was unrelated to trophic ecology. Differences in migration routes and stopovers may expose populations to distinct threats, and should be taken into consideration when defining units for conservation purposes and developing appropriate management strategies.

Lekking birds in a tropical forest forego sex for migration

Facultative, partially migratory animals provide a contemporary window into the evolution of migration, offering rare opportunities to examine the life-history trade-offs associated with migration. For the first time, to our knowledge, we describe the nature of these trade-offs, using a lek-breeding tropical bird, the white-ruffed manakin (Corapipo altera). Previous evidence indicated that weather drives post-breeding migration to lower elevations bringing condition-related benefits. Using elevation-sensitive stable isotope measurements and more than 1200 h of behavioural observations, we show that male manakins which migrate incur costs of diminished social status and matings with females the following breeding season. Because migratory tendency depends on inter-annual variation in weather, physical costs of displays and breeding prospects the following year, migratory decisions are subject to both natural and sexual selection, with the outcome of such decisions linked to changing climatic regimes.

Atlantic Leatherback Migratory Paths and Temporary Residence Areas

BackgroundSea turtles are long-distance migrants with considerable behavioural plasticity in terms of migratory patterns, habitat use and foraging sites within and among populations. However, for the most widely migrating turtle, the leatherback turtle Dermochelys coriacea, studies combining data from individuals of different populations are uncommon. Such studies are however critical to better understand intra- and inter-population variability and take it into account in the implementation of conservation strategies of this critically endangered species. Here, we investigated the movements and diving behaviour of 16 Atlantic leatherback turtles from three different nesting sites and one foraging site during their post-breeding migration to assess the potential determinants of intra- and inter-population variability in migratory patterns.Methodology/Principal FindingsUsing satellite-derived behavioural and oceanographic data, we show that turtles used Temporary Residence Areas (TRAs) distributed all around the Atlantic Ocean: 9 in the neritic domain and 13 in the oceanic domain. These TRAs did not share a common oceanographic determinant but on the contrary were associated with mesoscale surface oceanographic features of different types (i.e., altimetric features and/or surface chlorophyll a concentration). Conversely, turtles exhibited relatively similar horizontal and vertical behaviours when in TRAs (i.e., slow swimming velocity/sinuous path/shallow dives) suggesting foraging activity in these productive regions. Migratory paths and TRAs distribution showed interesting similarities with the trajectories of passive satellite-tracked drifters, suggesting that the general dispersion pattern of adults from the nesting sites may reflect the extent of passive dispersion initially experienced by hatchlings.Conclusions/SignificanceIntra- and inter-population behavioural variability may therefore be linked with initial hatchling drift scenarios and be highly influenced by environmental conditions. This high degree of behavioural plasticity in Atlantic leatherback turtles makes species-targeted conservation strategies challenging and stresses the need for a larger dataset (>100 individuals) for providing general recommendations in terms of conservation.

Post-breeding migration of four Long-tailed Skuas (Stercorarius longicaudus) from North and East Greenland to West Africa

The Long-tailed Skua (Stercorarius longicaudus) is a specialist predator of lemmings during the summer and hence an important component of the tundra ecosystems, but most of its life cycle takes place offshore and remains largely unknown outside of the breeding season. Using 9.5-g solar-powered satellite transmitters, we were able to document for the first time the post-breeding movements of the Long-tailed Skua, from its high-Arctic breeding-grounds in North and Eastern Greenland to the tropical waters of West Africa. The birds traveled the approximately 10,000 km of this migration in only 3–5 weeks, covering 800–900 km/day during active migration, which also occurred during nighttime. Leaving their breeding areas in August (except for one failed breeder), the Long-tailed Skuas first moved south along the coast of East Greenland towards a staging area off the Canadian Great Banks where they stayed for 1–3 weeks. From there, they crossed the Atlantic Ocean eastwards in just 1 week, entering African waters near the Madeira Archipelago in September. Although only four birds were monitored for 1.5–3 months, the data reveal that the migration routes between birds breeding in different locations and in different years were relatively similar.

Stress and parental care: Prolactin responses to acute stress throughout the breeding cycle in a long-lived bird

While the role of corticosterone in mediating the response of birds to acute stress is well established, it has recently been proposed that a decrease in prolactin levels following stress may complement corticosterone in redirecting resources away from breeding activities and towards behaviors promoting immediate survival. Here, for the first time, we detail changes in the prolactin stress response of birds throughout the breeding cycle. We then discuss the modulation of the corticosterone and prolactin stress responses over successive stages of breeding, differing in reproductive value and parental effort. In a long-lived Procellariiform seabird, the Manx shearwater Puffinus puffinus, we found that prolactin levels decreased in response to acute stress during incubation and mid chick-rearing but increased in response to stress during late chick-rearing and in non parenting birds, a pattern similar to that previously described for mammals. The high corticosterone stress response in pre-breeders was consistent with predictions based on reproductive value, but a similar response during late chick-rearing was not. This probably reflected foraging effort and a heightened importance of the parents’ own nutritional status at this stage of the season, in advance of post-breeding migration. We also found that baseline prolactin levels were maintained at high levels during chick-rearing and were only slightly lower during late chick-rearing and in failed breeders and non-breeders. These data suggest that prolactin may play a role in nestling care long beyond the brooding phase, that this is not due to birds spending long periods away from the colony and that prolactin secretion may be necessary for nest-guarding behavior.