PREY SELECTION AND TEMPORAL VARIATION IN THE DIET OF SUBANTARCTIC SKUAS AT INACCESSIBLE ISLAND,TRISTAN DA CUNHA

Ryan, P.G., & Moloney, C.L. 1991. Prey selection and temporal variation in the diet of Subantarctic Skuas at Inaccessible Island, Tristan da Cunha. Ostrich 62:52-58.

More than 2 500 prey items of Subantarctic Skuas Cuthuructa anturcricu were identified from regurgitated pellets collected at a roost of non-breeding skuas during summer 1989–90 at West Point, Inaccessible'Is-land. Most prey items (96,5%) were birds, primarily burrow-nesting procellariiforms. Sixteen bird species were recorfed in the diet, but five secies accounted for 94,6% of identifiable prey remains: Whitebellied Storm Petrel Fieettu grullariu (53,5%), Whitefaced Storm Petrel Pelugodrorna marina 15,1%), Broad-billed Prion Pacaptilu vittutu (14,0%), Great Shearwater Puffinus gruvis (7,3%), and Iommon Diving Petrel Pelecunoids urinutrix (4,7%). Petrel chicks were important in the diet, particularly Great Shearwaters. The main non-avian prey were fish and goose barnacles Lepas sp., but their importance may have been underestimated. The remains of a rat Rattus sp. presumably were derived from the neighbouring island of Tristan da Cunha; rats are not known to occur on Inaccessible Island. Prey size affected seabird availability to skuas, and dietary composition vaned throughout the summer in relation to seabird breeding cycles. Both these factors reduce the value of skua diet as an indicator of the relative abundances of burrow-nesting petrels. There was no correlation between skua diet and estimates of breeding bird abundance, but this may be a consequence of studying non-breeding skuas. Subantarctic Skua diet in 1989–90 was similar to that recorded in previous years, with consistent seasonal trends between years.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Impact of predation on king penguin Aptenodytes patagonicus in Crozet Archipelago

Predation can have major effects on population dynamics, but predator–prey interactions in marine ecosystems have rarely been studied. While the king penguin is one of the most studied seabirds, little is known about the impact of predation on its population dynamics. Here, we determine the impact of the main predators (giant petrels and skuas) on king penguin breeding success taking into account the nocturnal predation of petrels. We found that predation is the most important source of breeding failure for king penguins. The smallest chicks within crèches are the most hunted. The periphery of the colony suffers the highest risk of predation during summer. Our study shows the unequal quality of some areas inside the colony in terms of predation risk and breeding success, and points out the importance of timing in successful breeding.     

Predator‐avoidance behaviour in a nocturnal petrel exposed to a novel predator

Many species of bird recognize acoustic and visual cues given by their predators and have complex defence adaptations to reduce predation risk. Recognition of threats posed by specific predators and specialized anti‐predation behaviours are common. In this study we investigated predator recognition and anti‐predation behaviours in a pelagic seabird, Leach's Storm‐petrel Oceanodroma leucorhoa, at a site where predation risk from Great Skuas Stercorarius skua is exceptionally high. Leach's Storm‐petrels breed in burrows and come on land only at night. Counter‐predator adaptations were investigated correlatively in relation to changing natural light levels at night, and experimentally in relation to nocturnal visual and acoustic signals from Great Skuas. Colony attendance by Leach's Storm‐petrels was attuned to changes in light conditions at night and was highest when nights were darkest. This behaviour is likely to reduce predation risk on land; however, specific recognition of Great Skuas and specialized defence behaviours were not found. Leach's Storm‐petrels, in particular apparently non‐breeding individuals, were entirely naïve to the threat posed by Great Skuas and were captured easily in a variety of different ways, on the ground and in the air. Lack of specialized behavioural adaptations in Leach's Storm‐petrels against Great Skuas may be because spatial overlap of breeding distributions of these species appears to be a rare and recent phenomenon.     

SYSTEMATIC NOTES ON BIRDS OF THE CAPE PROVINCE: PART III.—PARISOMA SUBCAERULEUM (Vieillot): PART IV.—ANTHUS RICHARDI (Vieillot): PART V.—GALERIDA MAGNIROSTRIS Stephens

Fraser, M. W. 1984. Foods of Subantarctic Skuas on Inaccessible Island. Ostrich 55: 192–195.

Analysis of over 140 prey remains and 500 pellets of Subantarctic Skuas Catharacta antarctica collected from a site on Inaccessible Island. Tristan da Cunha, between October 1982 and January 1983 showed that 13 species of bird and one invertebrate were taken. Storm Petrels (Oceanitidae) made up 52% of the items recorded, Broadbilled Prions Pachyptila vittata 21%, and Common Diving Petrel Pelecanoides urinatrix 15%. Landbirds were rarely taken.     

PREDATION AND KLEPTOPARASITISM BY SKUAS IN A SHETLAND SEABIRD COLONY

Feeding methods and relations of Great Skuas and Arctic Skuas to prey were studied in a seabird colony at Hermaness, Shetland. Great Skuas obtained food by kleptoparasitism, predation and scavenging. They induced Gannets to regurgitate by interfering with their flight; grasping the Gannet by the wing or tail or pushing it down with the feet on its back. Gannets tried to escape by descending to the surface, and regurgitated during 12% of the chases, most frequently when pursued by several birds. Great Skuas caught Puffins by swooping at flocks in the colony. Puffins flying with fish to their young were also chased, releasing food on one fifth of the attacks, or escaping down to the sea and diving. Great Skuas also took Kittiwake nestlings by hovering and grasping the chick with the bill, killing and eating it on the surface. Adult Kittiwakes from nearby nests took to the air, mobbing the predator. More Kittiwakes were engaged in mobbing at unsuccessful than at successful predation attempts, indicating that colonial breeding may be of selective value under such predation. Two different estimates pointed to a Kittiwake nestling predation of 0–12 and 014 young per pair. Fledging success of Kittiwakes was estimated at 0–87-1-06 young per pair, considerably lower than at English colonies where predators are absent. In spite of the predation, the Kittiwake colony showed no signs of decrease. Agonistic behaviour and other evidence indicate that Great Skuas defend feeding territories at the seabird colony. Skuas, gulls and Fulmars competed for food at carcasses. Fulmars dominated and chased away skuas. Arctic Skuas deprived Puffins of food. They patrolled the cliff, intercepting Puffins arriving with fish, snatching it from their victim's bill, or inducing them to release fish. Puffins continuing their inward flight lost food more often (30%) than birds descending to the sea (15%)—sometimes diving below. This opportunity to escape may explain the lower success of skuas at Hermaness than at a Puffin colony farther inland from the shore (Grant 1971). Other factors being equal, proximity to the sea may thus reduce the risk of kleptoparasitism.     

Effects of predation risk on the nocturnal activity budgets of thin-billed prions Pachyptila belcheri on New Island, Falkland Islands

Predation is a major ecological and evolutionary driver of natural populations, greatly influencing fitness and behaviour of prey species. Small, long-lived petrels are vulnerable to predation at the breeding colonies and are expected to evolve behavioural strategies to minimize predation risks. Using an automatic nest monitoring system and nightly aerial counts, we examined the effect of vegetation cover and moonlight on colony attendance patterns and levels of burrow activity of breeding thin-billed prions, Pachyptila belcheri, on New Island, Falkland Islands. We further investigated how these parameters were related to predation by Falkland skuas. We monitored up to 32 nests in two habitats, one with Tussock grass and one with low vegetation cover. Individuals in both areas were more active at the nest before hatching, and those breeding in the low cover habitat were more active and arrived at the colony earlier, which might reflect an effect of reaction time over predation risk. Nocturnal activity peaks shifted in time as the season progressed, indicating behavioural adjustments to sunrise hours. Moon phase did not affect attendance and activity levels of breeders in either habitat or overall aerial activity, but influenced arrival time at the colony during chick-rearing, individuals arriving later in periods of full moon. Skua capture rates were positively correlated with aerial and nest activity but not with overall breeder attendance and were unaffected by moon phase. Thin-billed prions activity budgets are influenced by environmental parameters that affect their likelihood of being predated.     

Exploitation of inland-breeding Antarctic petrels by south polar skuas

During the austral summer of 1996/1997 we studied south polar skuas at Svarthamaren, Dronning Maud Land, Antarctica, where the world's largest known colony of Antarctic petrels is found. Our censuses suggested approximately 250 full-grown skuas and 140,000 breeding pairs of petrels were present. During their breeding season, skuas did not visit the open sea at least 200 km from the site; they relied entirely on prey caught and scavenged from the petrel colony. Because the site is so isolated, we asked whether the prey (petrels) had swamped the predators (skuas), or whether there was evidence that predator numbers were limited by the size of the prey population. Particularly at the end of the petrel incubation period, we found a close correspondence between the energy required by adult skuas and their chicks, ascertained from time budget studies, and the rate at which petrel eggs disappeared from the colony. This suggests that, in this closed system, the predator population was limited by the prey population, and that predator swamping was not an advantage that petrels gained by nesting in this remote location. Received: 12 April 1999 / Accepted: 30 June 1999     

Predation as a cost of sexual communication in nocturnal seabirds: an experimental approach using acoustic signals

Conspicuous behaviour, such as sexual advertisement, exposes animals to predation; mate attraction thus often conflicts with antipredator behaviour. We investigated whether an avian predator, the brown skua, Catharacta antarctica l?nnbergi, uses the mate attraction calls of colonial seabirds, the petrels. The majority of petrels attract mates at night and vocalizations are their main way of communicating. At our study sites, skua predation on nocturnal petrels was heavy, and concentrated particularly on a single species, the blue petrel, Halobaena caerulea. Using playback experiments, we showed that skuas can use male petrel calls as a cue for prey location and selection. This listening behaviour of skuas probably imposes a major constraint on advertising petrels, and especially on single males which face a trade-off between attracting females (which respond by calling in flight) and avoiding predation. We also investigated the consequences of this predation risk on the behaviour of petrels: a second set of playback experiments showed that the most heavily preyed on petrel species could use skua territorial calls to infer predation risk and stop calling thereafter, which may reduce conspicuousness and predation risk. Copyright 2000 The Association for the Study of Animal Behaviour.     

Nocturnal foraging by great skuas Stercorarius skua: implications for conservation of storm-petrel populations

At St Kilda, Outer Hebrides, a large colony of great skuas Stercorarius skua feed extensively on one of the largest colonies of Leach’s storm-petrels Oceanodroma leucorhoa in Europe, but little is known about the dynamics of this predator–prey system. Recently published population estimates of storm-petrels make it possible to estimate the impact of skua predation for the first time. Although skuas in the southern hemisphere catch petrels attending breeding colonies at night, it is not known whether congeners in the northern hemisphere also forage during the hours of darkness. We found (using radio-transmitters) that skuas regularly forage at night and (using light intensifying equipment) observed them catching storm-petrels at night. However, skuas also foraged during daylight hours, and it is unknown whether they might also catch storm-petrels at sea. Data on diet composition reveals that the proportion of storm-petrels in skua diet declined between 1996 and 1997, but remained constant thereafter. Although a large proportion of the storm-petrel prey is likely to consist of non-breeders, numbers consumed suggest that breeders and an unknown quantity of transients may also been eaten. The numbers of storm-petrels eaten are not sustainable and may result in substantial long-term population declines. Under current conditions, maintenance of large populations of both Leach’s storm-petrels and great skuas at St Kilda appears to be mutually exclusive.     

Hunting success of lions affected by the moon's phase in a wooded habitat

Variation in moonlight affects foraging, hunting and vigilance behaviours in many nocturnal species. Here, we explore the effect of moonlight on the movement and hunting behaviour of African lions. Previous studies found bright moonlight is associated with reduced hunting success; however, those studies were largely undertaken in open habitats where predators are easily seen by prey species on moonlit nights. In this study, we explored whether moonlight affected hunting behaviour and success in a largely wooded environment. Measures of short‐term lion movements (distance moved, displacement and path tortuosity) derived from GPS telemetry data were used as indicators of movement behaviour. Field observations of belly distension were used to assess recent food intake. Lions had greater belly distension (indicating feeding success) on dark nights. However, this change in feeding success was not reflected by lion movement patterns—there was no evidence that these changed across moon phases. There was no evidence that lions used more covered habitats on brighter nights to facilitate concealment.     

Patch use in time and space for a meso-predator in a risky world

Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.     

Determination of the absolute visual threshold of a nocturnal seabird, the Common Diving Petrel Pelecanoides urinatrix

Many of the smaller burrowing petrels are active at their colonies during the night. How they find their nesting burrows in the open terrain of an oceanic island or in the even more visually-taxing gloom under a forest canopy (Grubb 1974 and references therein) is a feat which has long intrigued naturalists (Lockley 1942). There is experimental evidence that vision is important in burrow location by the Manx Shearwater Puffinus puffinus (Brooke 1978, James 1986), while olfactory homing by Leach's Petrel Oceanodroma leucorhoa has also been suggested by Grubb (1974).
Additionally it seems likely that a number of petrel species catch a significant fraction of their food at night (Imber 1973, Prince & Francis 1984). Many of the squid presumed to be caught then are not bioluminescent (Imber 1973, Clarke et al. 1981), so their detection presumably demands good night vision. In the context of these observations, the dearth of information on the visual abilities of petrels is remarkable.
The present study investigated the absolute visual threshold of the Common Diving Petrel Pelecanoides urinatrix. Probably because the species is small (c. 130 g) and so vulnerable to predators such as Sub-antarctic Skuas Catharacta antarctica , it is strictly nocturnal at breeding colonies (Thoresen 1969, Payne & Prince 1979). It nests in burrows that are often located under tussock grass Poa spp. Whether the birds catch their crustacean prey (Payne & Prince 1979) by day or night is not known.     

Observations on the food of the south polar skua,Catharacta maccormicki near Davis,Antarctica

Summary Regurgitated boluses of undigested material were examined from six sites in East Antarctica. Skuas nesting near Adelie penguin, southern fulmar and Antarctic petrel colonies showed greater site fidelity in their feeding than did skuas nesting close to cape petrels or a Weddell seal pupping site.     

Predator–prey interactions between the South Polar skua Catharacta maccormicki and Antarctic tern Sterna vittata

Antarctic terns have to co‐exist in a limited space with their major nest predator, the skuas. We conducted artificial nest experiments to evaluate the roles of parental activity, nest location and nest and egg crypsis in this simple predator–prey system. Predation on artificial (inactive) nests was higher in traditional nesting sites than in sites previously not occupied by terns, which suggests that skuas memorized past tern breeding sites. Predation on artificial nests in inactive colonies was higher than in active (defended) colonies. Parental defense reduced predation in colonies to the level observed in artificial nests placed away from colonies. This suggests that communal defense can balance the costs of attracting predators to active colonies. Within colonies, predation was marginally higher on experimental eggs put in real nests than on bare ground. Although it seems that the presence of a nest is costly in terms of increased predation, reductions in nest size might be constrained by the need for protective nest structures and/or balanced by opposing selection on nest size. Predation did not differ markedly between artificial (quail) and real tern eggs. A simultaneous prey choice experiment showed that the observed predation rates reflected egg/nest detectability, rather than discrimination of egg types. In summary, nesting terns probably cannot avoid being detected, and they cannot defend their nest by attending them. Yet, by temporarily leaving the nest, they can defend it through communal predator mobbing, and at the same time, they can benefit from crypsis of unattended nest and eggs.     

Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Effect of predation by skuas on breeding success of the Cape petrel Daption capense at Nelson Island, Antarctica

Breeding success of Cape petrels at Nelson Island (South Shetland Islands) in 1991/1992 averaged 29%. Predation by skuas accounted for 76% of all nesting failures. Direct attacks prevailed over opportunistic predation. Intensity of observer disturbance alone could not account for the inter-colony variation in predation rate, which tended to increase with colony size, presence of nearby breeding skuas and local availability of other prey species (penguins). Predation was significantly lower on nests that could be attacked only, or more easily, from the air and nests with close neighbours. Selective predation on nests in which relatively narrower eggs were laid was independent of nest-site characteristics and laying date, suggesting an effect of parental age/experience on breeding success through antipredator behaviour. Development of oil-spitting behaviour in growing chicks balanced the parental attendance and effort in nest defence, which declined as chick age increased. Increased predation on lighter but not smaller chicks may reflect the lower ability of such chicks to defend themselves by spitting stomach oil. Seasonal variations of predation rate on Cape petrel nests did not result from predator swamping by numbers of simultaneously active nests, but rather reflected temporal availability of alternative prey. Success of all phases of nesting declined with later laying date. Received: 15 December 1997 / Accepted: 11 April 1998     

Origins, phytogeny and taxonomy of the gadfly petrels Pterodroma spp

Taxonomic characters of gadfly petrels (Procellariidae: Pterodroma spp.) are reviewed and the genus is redefined. The structure of the upper intestines, which have become helicoidally twisted to varying degrees in most species, is an important character not hitherto used. Results of a phylogenetic study of the genus based mainly on the development of helicoidal intestines agree substantially with findings from studies of the Mallophaga parasitizing these petrels. One species, the Kerguelen Petrel Lugensa brevirostris, previously classified in Pterodroma, is shown to have closer affinity with some fulmar genera. The Tahiti Petrel Pseudobulweria rostrata and its rare or extinct relatives, formerly placed in a superspecies of Pterodroma, are more closely related to the Procellaria and Bulweria petrels, and may be derived from the link between the subfamilies Procellariinae and Fulmarinae. From consideration of the fossil record, anatomical characters and Mallophaga, a phylogeny of the Procellariidae is proposed which supports its monophyletic origin. Gadfly petrels apparently descended from the fulmars through the ancestral lines of Snow Petrels Pagodroma, the Kerguelen Petrel and finally the Blue Petrel Halobaena, which may have given rise separately both to gadfly petrels and to the prions Pachyptila. A late Pliocene origin of Pterodroma in the vicinity of southern New Zealand is possible. The genus Pterodroma is divided into four subgenera, representing four connected radiations, and 29 species are recognised. In probable chronological order, Proaestrelata (new subgenus) contains five species and is restricted to the Pacific Ocean, Cookilaria comprises six species restricted as breeders to the South Pacific, Hallstroma has seven species mainly in the tropics of the Pacific, Indian and South Atlantic Oceans, whereas Pterodroma includes 11 species which extend as breeders from the southwest Pacific west to the South Atlantic and into the North Atlantic. Ecological, physical and physiological adaptations in the evolution of gadfly petrels are discussed.     

THE SIGNIFICANCE OF KLEPTOPARASITISM DURING THE BREEDING SEASON IN A COLONY OF ARCTIC SKUAS STERCORARIUS PARASITICUS IN ICELAND

Breeding and kleptoparasitism of Arctic Skuas was studied at Vik, Iceland, in 1973. Hatching success was 88.9%; fledging success was 0.27 fledglings per pair. The heavy chick mortality occurred mostly in the first week after hatching. In the early part of the breeding season skuas fed by robbing kittiwakes of their food at sea and by eating arthropods at the breeding grounds. At the time of hatching of skua eggs, which coincided with the hatching of Puffin eggs, skuas switched their feeding activities to kleptoparasitism of Puffins and fish so obtained was the principal item of most skuas' diet thereafter. In the first week post-hatching, the energy balance of an average adult skua pirating Puffins was estimated to be negative, but changed to positive a week later, although later, coinciding with an influx of non-breeders it turned back to negative. Arctic Terns which normally breed at Vik, and are exploited by skuas, failed to breed in 1973, and the abnormally heavy chick loss is therefore attributed to this failure of the terns. The success of the skuas kleptoparasitizing Puffins depends on the proximity of the interactions to the cliff or to the ocean, where Puffins seck refuge. Skuas catch fish mostly in the air, especially if it is dropped from high. Fish dropped from low is mostly taken by competing gulls and Ravens, which mostly control the ground and lower airspace. Skuas chasing in groups enhance their success, but the average success per member decreased with group increase. However, the success of the group-member in the ‘best’ position was equal to that of a single skua. Puffins carry 1 (large)-6 (smaller) fish, an approximately equal load irrespective of number of fish. Skuas preferentially chase Puffins carrying ‘large’ fish, thus maximizing their yield per effort. Arctic Skuas responded to changes in the numbers of arriving Puffins with a functional response, but their monitoring of the food supply was far from perfect.     

Yelkouan shearwater Puffinus yelkouan presence and behaviour at colonies: not only a moonlight question

We describe and test the influence of several environmental and biological factors on the presence and activity patterns of the Mediterranean endemic yelkouan shearwater Puffinus yelkouan at colonies. Bird arrival at breeding sites is highly correlated with nautical dusk for moonless or slightly moonlit nightfalls and correlated with moonset when the moon is visible. Breeding cycle and wind speed affect both the arrival times and presence at colonies. Bird activity also varies throughout the night and sex ratio on colonies throughout the breeding cycle in relation to breeding duties. Breeder and non-breeder behaviour particularly differs in the time spent on the ground outside burrows, without protection. Finally, factors other than moonlight can be essential in determining the presence and behaviour of petrels and shearwaters at breeding sites, and we need to determine how differences in behaviour at colonies could be related to differential predation risk.