Impacts of poor food availability on positive density dependence in a highly colonial seabird

For species with positive density dependence, costs and benefits of increasing density may depend on environmental conditions, but this has seldom been tested. By examining a colonial seabird (common guillemot) over a period of unprecedented poor food availability, we test two contrasting hypotheses suggesting that birds breeding at high density have: (i) greater leeway to increase foraging effort owing to more effective defence of unattended chicks against predators; and (ii) less leeway, owing to more attacks on unattended chicks by neighbouring adults. Supporting hypothesis 1, birds at high density increased provisioning rates and hence survival of chicks by foraging simultaneously with their partners, whereas at low density, unattended chicks were liable to be killed by predatory gulls and, unexpectedly, razorbills. Simultaneously, supporting hypothesis 2, heightened aggression towards unattended chicks at high density frequently resulted in infanticide, undermining benefits from collective defence against predators. Consequently, over 25 years, the magnitude of positive density dependence was independent of mean breeding success. These data indicate previously unsuspected trade-offs between costs and benefits of increasing density under changing environments. Previous generalizations about the importance of high density for reproductive success have so far remained robust, but such trade-offs could have unpredictable consequences for future population dynamics.     

Heat stress in a high-latitude seabird: effects of temperature and food supply on bathing and nest attendance of great skuas Catharacta skua

Birds such as great skuas Catharacta skua adapted for successful breeding at high latitudes may experience problems of heat dissipation in mild climates. Great skuas spend time bathing at freshwater sites close to breeding territories and here, we examine impacts of heat stress on bathing, foraging and nest attendance of adults during three breeding seasons with marked variation in the availability of prey (1-group sandeels Ammodytes marinus ). Adults exhibited diurnal variation in bathing activity that matched heat-stress conditions. Moreover more birds bathed on days of higher average heat stress, suggesting that bathing plays a role in thermoregulation. Bathing numbers were lower in years of poor food availability, when adult attendance at territories was low, probably because lower attendance reduced the opportunity for parents to bathe without leaving chicks unattended. Chicks are normally guarded by female parents and fed by males but under conditions of low food availability territorial attendance of breeding pairs was particularly low on days of high heat stress, with chicks regularly left unattended at air temperatures exceeding 14°C. Unattended chicks are at risk of being killed by neighbouring conspecifics and survival of chicks to fledging was low in the two years of low sandeel stocks. Our study indicates that for great skuas, indirect effects of climate change on prey stocks and direct effects on heat stress experienced by adults may be additive.     

Managing the Effects of Introduced Predators on Hawaiian Endangered Seabirds

Introduced predators are one of the greatest threats facing seabirds worldwide. We investigated the effects of multiple introduced predators on 2 endangered seabirds, the Newell's shearwater (Puffinus newelli) and the Hawaiian petrel (Pterodroma sandwichensis), on the island of Kauaʻi, Hawaiʻi, USA. Between 2011 and 2017, we recorded 309 depredations of which 35.6% were by feral cats, 50.2% by black rats (Rattus rattus), 10.4% by pigs (Sus scrofa; feral pigs), and 3.9% by barn owls (Tyto alba). Cats were the most destructive of the predators because they killed more breeding adults than chicks, which had repercussions on breeding probability in following years. Cats and rats were also the most prevalent of all the predators, depredating birds at all of the sites under consideration regardless of how remote or inaccessible. We also considered the effectiveness of predator control over the study period. Reproductive success at all sites increased once predator control operations were in place and depredations by all species except barn owls decreased. Furthermore, we modeled population trajectories for all sites with and without predator control. Without predator control, population trajectories at all sites declined rapidly over 50 years. With predator control operations in place, populations at all sites increased; thus, controlling introduced predators at endangered seabird colonies is important for their management. © 2020 The Wildlife Society.     

Parental attendance and squatting in the Kittiwake Rissa tridactyla during the rearing period

Parental attendance was studied in 1991 in the Cap Sizun Kittiwake colonies (Brittany, France). After a period of continuous guarding lasting on average 22 days, parents left their chicks unattended. Thereafter, parental attendance decreased regularly until fledging. The chick age when first left alone was on average 3 days lower for large broods than for single-chick broods. Moreover, whatever the brood size, chicks from late nests were younger when left unattended. Parental age affected the initiation of first absence. Younger parents reduced their attendance sooner than older parents. About 80% of the nests with chicks were visited by other adults at least once during the absence of the parents, and 50% were visited in the 3 days following the first absence of the breeders. These squatters were mainly failed breeders and prebreeders looking for a future breeding site. The results are discussed in terms of costs and benefits of chick neglect and comparisons were made with data from other studies in North Atlantic and Alaskan colonies. This reflected the flexibility of adult behaviour in relation to brood size and food availability.     

Populations and trends of Canadian Arctic seabirds

Canada’s eastern Arctic (Nunavut and Arctic Quebec—Nunavik, N of 60°) supports large numbers of seabirds in summer. Seabird breeding habitat in this region includes steep, rocky coasts and low-lying coasts backed by lowland sedge-meadow tundra. The former areas support colonial cliff- and scree-nesting seabirds, such as murres and fulmars; the latter inland or coastal seabirds, such as terns, gulls and jaegers. The region supports some 4 million breeding seabirds, of which the most numerous are thick-billed murres (Uria lomvia; 75%), black guillemots (Cepphus grylle; 9%), northern fulmars (Fulmarus glacialis; 8%) and black-legged kittiwakes (Rissa tridactyla; 6%). The majority of Arctic seabirds breed in a small number of very large colonies (>10,000 birds), but there are also substantial numbers of non-colonial or small-colony breeding populations that are scattered more widely (e.g. terns, guillemots). Population trends among Canadian Arctic seabirds over the past few decades have been variable, with no strongly negative trends except for the rare ivory gull (Pagophila eburnea): this contrasts with nearby Greenland, where several species have shown steep declines. Although current seabird trends raise only small cause for concern, climate amelioration may enable increased development activities in the north, potentially posing threats to some seabirds on their breeding grounds.     

Variance in prey abundance influences time budgets of breeding seabirds: evidence from pigeon guillemots Cepphus columba

We use data on pigeon guillemots Cepphus columba to test the hypothesis that discretionary time in breeding seabirds is correlated with variance in prey abundance. We measured the amount of time that guillemots spent at the colony before delivering fish to chicks ("resting time") in relation to fish abundance as measured by beach seines and bottom trawls. Radio telemetry showed that resting time was inversely correlated with time spent diving for fish during foraging trips (r=−0.95). Pigeon guillemots fed their chicks either Pacific sand lance Ammodytes hexapterus , a schooling midwater fish, which exhibited high interannual variance in abundance (CV=181%), or a variety of non-schooling demersal fishes, which were less variable in abundance (average CV=111%). Average resting times were 46% higher at colonies where schooling prey dominated the diet. Individuals at these colonies reduced resting times 32% during years of low food abundance, but did not reduce meal delivery rates. In contrast, individuals feeding on non-schooling fishes did not reduce resting times during low food years, but did reduce meal delivery rates by 27%. Interannual variance in resting times was greater for the schooling group than for the non-schooling group. We conclude from these differences that time allocation in pigeon guillemots is more flexible when variable schooling prey dominate diets. Resting times were also 27% lower for individuals feeding two-chick rather than one-chick broods. The combined effects of diet and brood size on adult time budgets may help to explain higher rates of brood reduction for pigeon guillemot chicks fed non-schooling fishes.     

Feeding frequency influences crèching age in the Dalmatian pelican, Pelecanus crispus

Crèching behaviour is common in colonial seabirds; nevertheless, the factors inducing chicks to aggregate remain relatively poorly understood. It has been proposed that brood size, laying date and nest attendance are important factors in the formation of a crèche. Moreover, in most species of pelicans, chicks join crèches following the development of homoeothermy and coincident with the end of the brooding behaviour. We studied effects of feeding rate, nest attendance, brood size, laying date and homoeothermy on the age at which chicks entered the crèche at a colony of Dalmatian pelicans (Pelecanus crispus), in Srebarna, Bulgaria. Single chicks were fed more frequently than chicks from two-chick broods. Unlike American white pelicans (Pelecanus erythrorhynchos), Dalmatian pelicans maintained brooding behaviour a further 9 days after chicks had developed thermoregulation abilities. In contrast to nests with two chicks, nests with only one chick were never left unattended by the parents before the chick reached the crèching stage. Laying date, nest attendance and brood size did not affect the age that the chick entered the crèche. The age the chick entered the crèche was not correlated with the age of homoeothermy acquisition, but chicks significantly joined the crèche at younger ages when the mean number of feeds per chick per day during the rearing period in the nest was higher. This result suggests an implication of growth rate in the crèching age. Joining the crèche earlier can provide benefits that could have strong implications for the chicks’ future reproductive lives.     

Re-Constructing Historical Adélie Penguin Abundance Estimates by Retrospectively Accounting for Detection Bias

Seabirds and other land-breeding marine predators are considered to be useful and practical indicators of the state of marine ecosystems because of their dependence on marine prey and the accessibility of their populations at breeding colonies. Historical counts of breeding populations of these higher-order marine predators are one of few data sources available for inferring past change in marine ecosystems. However, historical abundance estimates derived from these population counts may be subject to unrecognised bias and uncertainty because of variable attendance of birds at breeding colonies and variable timing of past population surveys. We retrospectively accounted for detection bias in historical abundance estimates of the colonial, land-breeding Adélie penguin through an analysis of 222 historical abundance estimates from 81 breeding sites in east Antarctica. The published abundance estimates were de-constructed to retrieve the raw count data and then re-constructed by applying contemporary adjustment factors obtained from remotely operating time-lapse cameras. The re-construction process incorporated spatial and temporal variation in phenology and attendance by using data from cameras deployed at multiple sites over multiple years and propagating this uncertainty through to the final revised abundance estimates. Our re-constructed abundance estimates were consistently higher and more uncertain than published estimates. The re-constructed estimates alter the conclusions reached for some sites in east Antarctica in recent assessments of long-term Adélie penguin population change. Our approach is applicable to abundance data for a wide range of colonial, land-breeding marine species including other penguin species, flying seabirds and marine mammals.     

Conflicts between touristic recreational activities and breeding shearwaters: short-term effect of artificial light and sound on chick weight

Human disturbances are increasingly becoming a conservation concern for many populations of colonial seabirds. Colonially reproducing species are particularly vulnerable to localised disturbances because detrimental elements can simultaneously affect the entire population. Studies of petrels and shearwaters have shown that light pollution, in particular, can be harmful for both fledglings and adults, but little is known of the way such anthropogenic elements affect the quality of parental care at the nest. Chick provisioning in petrels and shearwaters occurs exclusively at night and is also negatively correlated with the amount of moonlight. We tested the hypothesis that high-intensity light and sound disturbances will disrupt nest attendance and thus affect weight gain in chicks but that the magnitude of such effects would be modulated by moonlight conditions. We measured the effect of two outdoor disco events on overnight weight gain in 26 chicks of Scopoli’s shearwaters (Calonectris diomedea) from a breeding colony on Linosa Island. The two disco events occurred under contrasting moonlight conditions (moonless vs moonlight). Chicks situated closer to the disturbance gained significantly less weight compared to conspecifics from nests further away but the effect was only evident on the moonless night.Our results suggest that light and sound disturbances can have a negative effect on parental care in C. diomedea but moonlight might moderate the bird’s perception and thus the magnitude of the disturbance. However, while occasional disturbances may impact short-term weight gain in C. diomedea chicks, such effects are not perceivable at fledging when measured as differences in the weight or the date at which they left the nest.     

Predation by Ravens Corvus corax on Brunnich's Guillemot Uria lomvia eggs and chicks and its possible impact on breeding site selection

We studied the behaviour of Ravens Corvus corax taking the eggs or chicks of Brunnich's Guillemots Uria lomvia at East Digges Island, Northwest Territories, Canada. We classified the characteristics of the breeding sites attacked by Ravens according to their location on the cliffs, the width of the breeding ledge, the number of adjacent rock faces and the density of nearby breeding guillemots. Ravens selectively attacked peripheral sites near the top of the occupied area, those on narrow ledges and those with few close neighbours. The risk of predation at the most vulnerable sites was more than seven times the average. For the most part, breeding success at 419 sites near the cliff top conformed to predictions based on the observations of Raven predation. However, losses observed over 3 years suggested that more eggs fall accidentally than are taken by avian predators. We concluded that the risks of predation and of accidental dislodgement both contribute to choice of nest sites by Brunnich's Guillemots. The effect of multiple close neighbours in deterring predation by Ravens may be important in determining the density of guillemot breeding sites within the colony.     

Sex‐specific lateralization during aggressive interactions in breeding king penguins

Brain and behavioral asymmetries (termed "lateralization"; e.g., preferential eye‐use) have been mostly described in controlled laboratory conditions, although striking similarities of hemispheric brain control for specific behaviors have also been shown in the wild. Visual lateralization may provide ecological advantages by allowing complementary roles played by the left–right lateral and frontal visual field in distant or close motion detection of predators or other threats. In this study, we tested for lateralization in aggressive behavior in wild king penguins (Aptenodytes patagonicus), seabirds breeding in a context of strong colonial aggressiveness, and subject to on‐land‐based predation of their egg or chick. We show that males initiated more agonistic interactions when a congener was located in their right frontal visual field and in their left lateral visual field. The results obtained in females were the exact opposite for each subdivision of their visual fields. Complementary lateralization in male and female penguins may be part of a more general phenomenon, allowing partners to coordinate their behavior during reproduction. This may be especially true during the period of courtship, during which these seasonally monogamous and monomorphic seabirds engage in mutual mate choice based on a complex and ritualized display of ornaments located on the left and right lateral sides of the head. Those results open exciting questions as to whether hemispheric control of aggression is a commonly selected phenotypic trait across colonial seabirds.     

Benefits and costs of rapid growth in common murre chicks Uria aalge

While accelerated growth can be advantageous to nestling birds, there may be a tradeoff between rapid growth and resistance to food shortages. Common murres Uria aalg e are colonial seabirds that benefit from reproductive synchrony. Individuals that lay eggs late should benefit if they produce chicks capable of growing quickly and fledging synchronously with their neighbors. In this study, we controlled food provisioning of captive-hatched common murre chicks from a single subcolony and examined differences in growth between early-hatched individuals and their later-hatched neighbors. We assessed potential costs of rapid growth by comparing growth of chicks fed ad libitum with their growth under food restricted conditions. Chicks that hatched later were heavier, ate more and gained body mass more quickly than chicks that hatched earlier. Late-hatched chicks grew quickly enough to reach the same mass as their early-hatched neighbors in five days. However, chicks that grew more quickly under ad libitum food conditions grew more slowly when food was restricted. We conclude that murres that lay eggs late may synchronize their reproduction with early-laying neighbors by producing rapidly growing chicks. However, the ability to compensate for late hatching by growing quickly can be costly when food becomes limited.     

Prospecting enhances breeding success of first-time breeders in the great cormorant, Phalacrocorax carbo sinensis

In many species of colonial seabirds, young birds visit colonies in the years before they start breeding. This prospecting behaviour may allow them to obtain information that could enhance their future breeding success. We examined the reproductive consequences of prospecting behaviour in the colonial great cormorant, and found support for this idea. New breeders that had been prospecting actively in the previous year obtained breeding sites of higher quality (i.e. closer to sites where conspecifics had fledged young in the previous year) and had higher breeding success than those that had been less active. Prospecting occurred mostly late in the breeding season, and coincided with the time when the majority of the eggs had hatched but before the chicks started fledging, that is, when breeding success in the colony reflected habitat suitability. These results are thus consistent with the use of conspecific reproductive performance as a cue for the quality of a breeding habitat as expected from the 'performance-based conspecific attraction hypothesis'. Copyright 1999 The Association for the Study of Animal Behaviour.     

Sex-biased disruptive behaviour in breeding crested penguins

Colonial breeding is common in seabirds, and may provide individuals with benefits such as increased protection from predators by joint defence, improved information exchange and enhanced access to mates. However, the presence of large numbers of individuals in breeding colonies may also lead to interference, especially where conspecific behaviour disrupts the normal chick-rearing routine. Using standardised video recordings, we describe and quantify for the first time such disruptive behaviour by conspecifics in penguins. This disruptive behaviour appeared to be common in chick-feeding Southern rockhopper penguins Eudyptes chrysocome and its occurrence did not differ significantly among the colony areas where observations were made. Females were commonly (77 % of 111 observation sessions) disrupted by other adults when attempting to feed chicks alone during the crèche stage. They consequently reduced the time spent with their chicks. Disrupted chicks compensated by begging more intensely when their female parents were present, and they did not eat significantly less often than undisrupted chicks. In contrast, disruptive behaviour towards chick-feeding males or pairs was never observed. This study supplements knowledge about a disruptive behaviour that appears to be common but for which the function remains unclear in penguins, as the intruders did not appear to gain any visible benefits.     

Breeding synchrony and predator specialization: A test of the predator swamping hypothesis in seabirds

Reproductive synchrony is a widespread phenomenon that is predicted to be adaptive for prey with specialist predators but not for those with generalist ones. I tested this prediction in three polar seabird species characterized by different levels of predator specialization. In the Antarctic petrel, for which the only predator was highly specialized, hatching dates were highly synchronous and chicks that hatched close to the mean hatching date had a higher survival. In black‐legged kittiwakes and Brünnich's guillemots, whose predators were generalists, breeding was less synchronous and there was no fitness advantage in hatching close to the mean. This study emphasizes the potential importance of the relative timing of reproduction for individual fitness and supports the hypothesis that the adaptive value of breeding synchrony depends on the predator functional response.     

Investigating the effects of age‐related spatial structuring on the transmission of a tick‐borne virus in a colonially breeding host

Higher pathogen and parasite transmission is considered a universal cost of colonial breeding due to the physical proximity of colony members. However, this has rarely been tested in natural colonies, which are structured entities, whose members interact with a subset of individuals and differ in their infection histories. We use a population of common guillemots, Uria aalge, infected by a tick‐borne virus, Great Island virus, to explore how age‐related spatial structuring can influence the infection costs borne by different members of a breeding colony. Previous work has shown that the per‐susceptible risk of infection (force of infection) is different for prebreeding (immature) and breeding (adult) guillemots which occupy different areas of the colony. We developed a mathematical model which showed that this difference in infection risk can only be maintained if mixing between these age groups is low. To estimate mixing between age groups, we recorded the movements of 63 individually recognizable, prebreeding guillemots in four different parts of a major colony in the North Sea during the breeding season. Prebreeding guillemots infrequently entered breeding areas (in only 26% of watches), though with marked differences in frequency of entry among individuals and more entries toward the end of the breeding season. Once entered, the proportion of time spent in breeding areas by prebreeding guillemots also varied between different parts of the colony. Our data and model predictions indicate low levels of age‐group mixing, limiting exposure of breeding guillemots to infection. However, they also suggest that prebreeding guillemots have the potential to play an important role in driving infection dynamics. This highlights the sensitivity of breeding colonies to changes in the behavior of their members—a subject of particular importance in the context of global environmental change.     

Evidence that tufted puffins Fratercula cirrhata use colony overflights to reduce kleptoparasitism risk

Predation, foraging and mating costs are critical factors shaping life histories. Among colonial seabirds, colony overflights may enhance foraging or mating success, or diminish the risk of predation and kleptoparasitism. The latter possibility is difficult to test because low predation or kleptoparasitism rates could be due either to low danger or to effective counter-tactics by prey. Tufted puffins Fratercula cirrhata breeding at a large colony in British Columbia, Canada, deliver several loads of fish each day to their nestlings and are targets for kleptoparasitism by glaucous-winged gulls Larus glaucescens . In the present study, we documented the ecological conditions under which overflights occurred in order to assess when overflights were made and to statistically isolate the effect of overflights on kleptoparasitism risk at this site. Load-carrying puffins engaged in overflights under ecological conditions associated with relatively high rates of kleptoparasitism. Further, when ecological factors determining risk were statistically controlled, overflights were correlated with marginally lower chances of kleptoparasitism than when the risk factors were ignored. The results suggest that breeding puffins at this site use overflights for kleptoparasite avoidance. This tactic is used sparingly, suggesting it is costly. Costs of overflight behaviour might contribute to the impact of kleptoparasitism on the breeding success of tufted puffins.     

Validating growth and development of a seabird as an indicator of food availability: captive‐reared Caspian Tern chicks fed ad libitum and restricted diets

ABSTRACT For seabirds raising young under conditions of limited food availability, reducing chick provisioning and chick growth rates are the primary means available to avoid abandonment of a breeding effort. For most seabirds, however, baseline data characterizing chick growth and development under known feeding conditions are unavailable, so it is difficult to evaluate chick nutritional status as it relates to foraging conditions near breeding colonies. To address this need, we examined the growth and development of young Caspian Terns (Hydroprogne caspia), a cosmopolitan, generalist piscivore, reared in captivity and fed ad libitum and restricted (ca. one‐third lower caloric intake) diets. Ad libitum‐fed chicks grew at similar rates and achieved a similar size at fledging as previously documented for chicks in the wild and had energetic demands that closely matched allometric predictions. We identified three general characteristics of food‐restricted Caspian Tern chicks compared to ad libitum chicks: (1) lower age‐specific body mass, (2) lower age‐specific skeletal and feather size, such as wing chord length, and (3) heightened levels of corticosterone in blood, both for baseline levels and in response to acute stress. Effects of diet restriction on feather growth (10–11% slower growth in diet‐restricted chicks) were less pronounced than effects on structural growth (37–52% slower growth) and body mass (24% lower at fledging age), apparently due to preferential allocation of food resources to maintain plumage growth. Our results suggest that measurements of chick body mass and feather development (e.g., wing chord or primary length) or measurement of corticosterone levels in the blood would allow useful evaluation of the nutritional status of chicks reared in the wild and of food availability in the foraging range of adults. Such evaluations could also inform demography studies (e.g., predict future recruitment) and assist in evaluating designated piscivorous waterbird conservation (colony) sites.     

Aggression by adult South American Terns toward conspecific chicks

ABSTRACT Infanticide and nonfatal aggression by adults toward unfamiliar chicks have been widely reported in colonial birds, and can be an important cause of chick mortality. We studied intraspecific aggression by adults toward chicks at a South American Tern (Sterna hirundinacea) colony in Patagonia, Argentina, during 2005 to characterize this behavior, evaluate its relationship with nesting density, chick age and microhabitat characteristics, and assess its effect on breeding success. Of 111 chicks in the study area, 45% were attacked at least once. Chicks older than 9 d posthatching were more likely to be attacked than younger chicks, and unattended chicks were more likely to be attacked than guarded chicks (88 vs. 12%). Chicks were also attacked more often when in their own territories (76% of cases), but were less likely to be attacked in territories with more vegetation cover and high‐quality shelters (i.e., vegetation with characteristics that prevented adults from reaching chicks). The number of aggression events was not related to nest density. At least 8% of the chicks in our study area died as a result of adult intraspecific aggression. Our results indicate that intraspecific aggression by adult South American Terns toward chicks is relatively common in the Punta Loma colony and should not be underestimated as a factor affecting their breeding success.