Foraging strategies of incubating and brooding king penguins Aptenodytes patagonicus

For oceanic birds like king penguins, a major constraint is the separation of foraging areas from the breeding colony, largely because swimming increases foraging costs. However, the relationship between foraging strategy and breeding stage has been poorly investigated. Using time-depth recorders, we studied the diving behaviour of two groups of king penguins that were either incubating or brooding chicks at Crozet Islands (Southern Indian Ocean) at the same period of the year. Although birds with chicks had the highest predicted energy demand, they made foraging trips half as long as incubating birds (6 vs. 14 days) and modified their time and depth utilisation. Birds with chicks dived deeper during daylight (mean maximum depth of 280 m vs. 205 m for those incubating). At night, birds with chicks spent twice as much time diving as those incubating, but birds at both stages never dived beyond 30 m. Movements to greater depths by brooding birds are consistent with the vertical distribution of myctophid fish which are the main prey. As chick provisioning limits trip duration, it is suggested that it is more efficient for parents to change their diving patterns rather than to restrict their foraging range. Received: 23 June 1997 / Accepted: 3 November 1997     

AIRCRAFT SOUND AND DISTURBANCE TO BOWHEAD AND BELUGA WHALES DURING SPRING MIGRATION IN THE ALASKAN BEAUFORT SEA

Short‐term behavioral responses of bowhead whales (Balaena mysticetus) and beluga whales (Delphinapterus leucas) to a Bell 212 helicopter and Twin Otter fixed‐wing aircraft were observed opportunistically during four spring seasons (1989–1991 and 1994). Behaviors classified as reactions consisted of short surfacings, immediate dives or turns, changes in behavior state, vigorous swimming, and breaching. The helicopter elicited fewer detectable responses by bowheads (14% of 63 groups) than by belugas (38% of 40). Most observed reactions by bowheads (63%) and belugas (86%) occurred when the helicopter was at altitudes ≤150 m and lateral distances ≤250 m. Belugas reacted significantly more frequently during overflights at lateral distances ≤250 m than at longer lateral distances (P= 0.004). When the helicopter was on the ice with engines running, 7 of 14 groups of belugas reacted, up to 320 m away, sometimes with small‐scale (≤100 m) diversion; only 1 of 8 groups of bowheads reacted. For the fixed‐wing aircraft, few bowheads (2.2%) or belugas (3.2%) were observed to react to overflights at altitudes 60–460 m. Most observed reactions by bowheads (73%) and belugas (70%) occurred when the fixed‐wing aircraft was at altitudes ≤182 m and lateral distances ≤250 m. However, the proportions reacting, especially to low‐altitude flights (e. g., ≤182 m), were underestimated for both species because observation opportunities were brief. Even so, reactions were more common when the aircraft was low (≤182 m): P= 0.009 for belugas, P= 0.06 for bowheads. There was little if any reaction by bowheads when the aircraft circled at altitude 460 m and radius 1 km. Aircraft sounds measured underwater at depths 3 m and 18 m showed that a Bell 212 helicopter was 7–17.5 dB noisier than a Twin Otter (10–500 Hz band). Bell 212 sound consisted mainly of main rotor tones ahead of the helicopter and tail rotor tones behind it. Twin Otter sound contained fewer prominent tones. Peak sound level as received underwater was inversely related to aircraft altitude, and received levels at 3 m depth averaged 2.5 dB higher than at 18 m depth. The dominant low‐frequency components of aircraft sound are presumed to be readily audible to bowheads. For belugas, these components may be inaudible, or at most only weakly audible. Mid‐frequency sound components, visual cues, or both, are probably important in eliciting beluga reactions to aircraft.     

Disturbance of emperor penguin Aptenodytes forsteri chicks by helicopters

Creching emperor penguin (Aptenodytes forsteri) chicks were exposed to two overflights by a Sikorsky S-76, twin engine helicopter at 1000 m (3300 ft), a current operational guideline of the Australian Antarctic Division for helicopter activity in Antarctica. The flights were conducted on the same day but under different wind conditions: a morning flight with a 10-knot (18 km h⁻¹) katabatic wind blowing perpendicular to the direction of helicopter travel, and an afternoon flight with virtually no wind. Background noise levels recorded in the morning before the helicopter flight were significantly higher than in the afternoon, but these differences were not detectable when the helicopter was overhead. There were also no significant differences in the way chicks responded to helicopters between the morning and afternoon flight. All chicks became more vigilant when the helicopter approached and 69% either walked or ran, generally moving less than 10 m toward other chicks (i.e. not scattering). Most chicks (83%) displayed flipper-flapping, probably indicating nervous apprehension. This behaviour was seldom displayed in the absence of disturbance. Although all effects were relatively transitory, the results support the introduction of a more conservative guideline of 1500 m (5000 ft) minimum overflight altitude for helicopter operations around breeding localities of this species. Accepted: 31 May 1999     

Stomach stones in king penguin chicks

Many animals that possess a gizzard swallow stones or sandy grit, supposedly to aid in the mechanical breakdown of food. While this has been well documented in the literature, our study is the first to report the presence of stones in the gizzard of king penguin chicks. We found stones, so called ‘gastroliths’, in the pyloric region of the gizzard, the part of the digestive tract that is specialised for the mechanical breakdown of food. Stones were already present in the gizzard of chicks and, hence, during the first year of the life of king penguins, which is spent on land. Some chicks were found to have more than 130 stones (0.5–22 mm in size) in their gizzard. The gastroliths we found in king penguins are of the same geological origin as rocks present at the colony, which suggests that birds swallowed them there. The functional role of gastroliths in penguin chicks and adults is still unknown. We discuss the potential roles that these gastroliths might play in king penguins (i.e. aid in digestion, buoyancy control during foraging at sea, adaptation to fasting).     

Krill-feeding behaviour of gentoo penguins as shown by animal-borne camera loggers

Animal-borne camera loggers were used to examine the patterns of prey encounter and feeding behaviour of gentoo penguins at King George Island, Antarctica. The still images from the camera loggers showed that the penguins encountered the swarms of krill for 25.5% (range: 8–38%) of their dives (>5 m) on average, during their foraging trips (mean duration of 5.4 h, n = 7 trips). They encountered krill swarms during the dives to 10–70 m depth, in pelagic as well as benthic habitats. In the benthic habitat, the penguins swam just above the sea floor and headed downward over a krill swarm, probably using the sea floor to assist them to feed on mobile swarms. The shallow coastal waters would be the important foraging habitat of gentoo penguins breeding in King George Island.     

Is Penguin banding harmless?

Thousands of penguins are banded annually world-wide, even though little is known about the potential impact of these flipper bands. In this paper, the possible effect of banding on the survivorship, breeding frequency and other ecological factors on king penguins was investigated. The extended laying period (3–4 months) of the king penguin allows the observation of non-lethal effects that could influence the time of laying and thus the reproductive success. Three hundred and eighty-three breeding king penguins on the colony of “la Baie du Marin” on Possession Island, Crozet Archipelago were either single- or double-flipper banded. The results show that the returning birds were laying late the following breeding season, and that double-banded birds lay significantly later than single-banded birds. Furthermore, our data suggest a lower return rate for double-banded birds than for single-banded birds (45% as opposed to 76%). The low return rate of single-banded birds, when compared to an interannual survivorship estimated to be 96.5%, also raises questions concerning the potential impact of single banding. Received: 2 January 1998 / Accepted: 23 May 1998     

Comparison of behaviour,body mass,haematocrit level,site fidelity and survival between infested and non-infested king penguin<Emphasis Type="Italic"> Aptenodytes patagonicus</Emphasis> by ticks<Emphasis Type="Italic"> Ixodes uriae</Emphasis>

The tick Ixodes uriae is widespread in subantarctic areas. This tick species parasitises a large number of seabird species and may affect their population dynamics. Adult king penguins are significant hosts for I. uriae in Crozet Archipelago. We compared the body mass, haematocrit level and behaviour during incubation according to tick infestation. Tick infestation had no effect on the body mass, haematocrit or behaviour of incubating penguins. Contrary to what might have been expected, birds breeding in the infested area did not show a lower fidelity to that area the next breeding season in comparison to birds breeding in the non-infested area. Survival was not different between the two groups over the following 32 months.     

Aggressiveness in king penguins in relation to reproductive status and territory location

King penguins, Aptenodytes patagonicus, vigorously defend small territories in very dense colonies. The egg-laying season lasts approximately 4 months, but only pairs that reproduce during the first half of the period succeed in fledging a chick. I examined various factors affecting aggressiveness of king penguins during the breeding season and focused on the differences between central and peripheral territories. Pairs on peripheral territories experienced twice as many encounters with avian predators as did central birds. The vast majority of peripheral birds were late breeders, indicating that reproductive success was very low among penguins defending a territory on the edge of the colony. Time invested in territory defence and rate of agonistic encounters between breeding neighbours increased from the incubation to the brooding period. Parents gave most threat displays to territorial neighbours when the chick was very young and just before crèche formation. Distance to colony edge was not related to aggressiveness in incubating birds, however, the rate of pecking and flipper blows increased from the edge to the centre during brooding. In addition, aggressiveness of breeding penguins towards travelling birds trespassing into their territory increased with distance to edge. Early breeders were not more aggressive than late breeders but the proportion of time spent in territory defence increased with the number of days a bird had spent incubating. As expected, I did not detect any sex difference in aggressive behaviour. Birds occupying territories on the beach were generally more aggressive during the incubation period than those located on the valley sides. Reproductive status (incubating versus brooding) and territory location were the main factors explaining the various levels of aggressiveness observed in breeding king penguins. Copyright 2000 The Association for the Study of Animal Behaviour.     

The Apache Longbow–Hellfire Missile Test at Yuma Proving Ground: Introduction and Problem Formulation for a Multiple Stressor Risk Assessment

An ecological risk assessment was conducted at Yuma Proving Ground, Arizona, as a demonstration of the Military Ecological Risk Assessment Framework (MERAF). The focus of the assessment was a testing program at the Cibola Range, which involved an Apache Longbow helicopter firing Hellfire missiles at moving targets, that is, M60-A1 tanks. The problem formulation for the assessment included conceptual models for three component activities of the test, helicopter overflight, missile firing, and tracked vehicle movement, and two ecological endpoint entities, woody desert wash communities and desert mule deer (Odocoileus hemionus crooki) populations. An activity-specific risk assessment framework was available to provide guidance for assessing risks associated with aircraft overflights. Key environmental features of the assessment area include barren desert pavement and tree-lined desert washes. The primary stressors associated with helicopter overflights were sound and the view of the aircraft. The primary stressor associated with Hellfire missile firing was sound. The principal stressor associated with tracked vehicle movement was soil disturbance, and a resulting, secondary stressor was hydrological change. Water loss to desert washes and wash vegetation was expected to result from increased ponding, infiltration, and/or evaporation associated with disturbances to desert pavement. A plan for estimating integrated risks from the three military activities was included in the problem formulation.     

Diving patterns of female macaroni penguins breeding on Marion Island, South Africa

Despite the large biomass of macaroni penguins Eudyptes chrysolophus in the Southern Ocean, their feeding ecology is poorly known at some important breeding localities. We investigated the diving behaviour and diet of female macaroni penguins feeding small chicks on Marion Island (46o52′S, 37o5′E), South Africa, one of the species’ most northerly breeding sites, supporting 4% of their global population. We then compared our results with similar studies from other localities. In December 2008, we collected information on 12 foraging trips from 6 individuals using time-depth recorders, as well as diet from 42 individuals. Median trip duration was 22.8 h (5.6–80.8 h). Penguins performed 42.8 ± 15.9 dives per hour at sea, with dive depths averaging 24.6 ± 8.6 m and lasting 40.8 ± 12.1 s, although 74.3% of dives were <10 m. Euphasids dominated their diet (86% by mass), mainly Thysanoessa vicina. A second peak in dive depths at 55–80 m might reflect the 12% of fish in their diet. The substantial proportion of shallow night dives (30% of total dives) suggests some foraging occurs at night. Differences in diving patterns of individual macaroni penguins in this study confirmed the behavioural flexibility of these birds reported from other breeding localities. However, most other studies assumed that dives <3–5 m were commuting dives whereas our study suggests that at least some prey are caught during shallow dives. We highlight how different analytical methods can change the outcome of studies. Despite macaroni penguins’ apparent flexibility in foraging behaviour during the breeding season, their numbers are decreasing globally. Further investigations of their foraging behaviour are needed to assess potential competition with other predators and krill fisheries.     

Behavioral response of polar bears to aircraft activity on the northern coast of Alaska

The rapid loss of arctic sea ice is forcing a larger proportion of the Southern Beaufort Sea polar bear (Ursus maritimus) population to spend more time on land, increasing chances of negative interactions between people and bears. In the United States, the Marine Mammal Protection Act (MMPA) protects polar bears from incidental disturbance from human activities. For the remote and roadless areas of northern Alaska, USA, effective management of small aircraft activity is necessary to limit disturbance, but effects of overflights on polar bear behavior are largely unknown. During 2021 and 2022, we intentionally exposed polar bears (n = 115) to systematic aircraft activity (helicopter, fixed-wing) until we observed a disruption of behavior that qualified as a level B take response (e.g., abrupt change in activity or movement) under the MMPA. We used a Bayesian logistic regression to determine what factors influence and can be used to predict when a polar bear will exhibit a level B take response and estimate the probability of an aircraft eliciting a level B take response at different altitudes above the polar bear. Aircraft type, flight altitude, landscape (barrier islands vs. mainland), and bear behavior (active vs. inactive) upon initial aircraft encounter were all important predictors of take. Probability of take rapidly increased with a decrease in flight altitude starting at 450 m for helicopter and 300 m for fixed-wing aircraft. Active (e.g., standing, walking) polar bears on barrier-island landscapes were more likely to experience take than inactive (e.g., bedded) bears on mainland landscapes. Our findings can help with assessments and management plans by quantifying disturbance to polar bears from current and future human activity that involves aircraft use.     

Seasonal change in foraging areas and dive depths of breeding king penguins at Heard Island

The seasonal variation in the foraging behaviour of king penguins (Aptenodytes patagonicus) was studied at Heard Island (53°05′S, 73°30′E) during 1992/1993. On seven occasions throughout the breeding cycle, time-depth-light recorders were deployed on breeding adults to record the dive activities and foraging. Foraging locations changed with season: in autumn and spring 1992, adults foraged between 48–52°S and 74–78°E, about 370 km NNE of Heard Island close to the Polar Front. Two penguins tracked in winter travelled 2220 km east of Heard Island (95°E) along the northern ice limit, and 1220 km south of Heard Island to approximately 65°S, respectively. In spring (October), the penguins again foraged further north than during winter. The foraging area utilised in October overlapped the area where the penguins foraged in March/April. The penguins' diving behaviour also varied seasonally: the modal depth of deep dives (>50 m) increased from about 100 m in February to 220 m in October. Mean dive depths increased from 70 ± 52 m in March 1992 to 160 ± 68 m in August 1992. Penguins dived deep (>50 m) only during daylight hours (16 h in February, 9 h in July). Mean dive durations ranged from 2.9 ± 1.1 min in March 1992 to 5.1 ± 1.2 min in August 1992. Associated with changes in foraging location and dive behaviour was a change in diet composition: during summer the penguins ingested mainly myctophid fish (>90%) while in winter the most important diet item was squid. Accepted: 19 October 1998     

Post-breeding dispersal of Ad��lie penguins (Pygoscelis adeliae) nesting at Signy Island, South Orkney Islands

ARGOS satellite telemetry and Global Location Sensors (geolocators) were used to identify the moult locations and the winter foraging dispersal of Adélie penguins after they left their breeding colonies on Signy Island in the South Orkney Islands. Animals were tracked during the period December 2004 to October 2005. All birds displayed a similar pattern of migratory behaviour, remaining away from colonies for approximately 9 months, at distances of up to 2,235 km. Moult locations were within the pack ice. Mean daily travel speeds to the moult locations were significantly faster when moving through open water than through pack ice. Moult occurred during February/March within a narrow latitudinal range (65–71°S), at a mean distance of 126 km from the ice edge; the mean duration of individual moult was c. 18.6 days. After moult, penguins spent the subsequent winter months moving north or north-eastward within the expanding winter pack ice, at a mean distance of 216 km from the ice edge, and in areas with ice cover >80%. The penguins returned to the vicinity of their colony between September 26 and October 22, 2005. This dependence of Adélie penguins on sea ice habitat suggests that any further reductions in sea ice extent in the Weddell Sea region would potentially have important impacts on the population processes of this pagophilic species.     

Vulture flight behavior and implications for aircraft safety

Growing vulture populations represent increasing hazards to civil and military aircraft. To assess vulture flight behavior and activity patterns at the Marine Corps Air Station in Beaufort, South Carolina, we equipped 11 black vultures (Coragyps atratus) and 11 turkey vultures (Cathartes aura) with solar-powered Global Positioning System (GPS) satellite transmitters during a 2-year study (1 Oct 2006–30 Sep 2008). Turkey vultures had larger seasonal home ranges than did black vultures, and 2 turkey vultures made round-trips to Florida. Black vultures consistently spent less time in flight (8.4%) than did turkey vultures (18.9%), and black vultures flew at higher altitudes than did turkey vultures in all seasons except summer when altitudinal distributions (above ground level) did not differ. Although we recorded maximum altitudes of 1,578 m for black vultures and 1,378 for turkey vultures, most flights were low altitude. A matrix of vulture flight altitude versus time of day revealed that >60% of vulture flight activity occurred from 4 hr to 9 hr after sunrise at altitudes below 200 m. Continuation of aggressive harassment coupled with flexible training schedules to avoid times and altitudes of high vulture activity will decrease hazards to aircraft posed by these birds. © 2011 The Wildlife Society.     

Comparison of foraging strategies of incubating king penguins Aptenodytes patagonicus from Macquarie and Heard islands

The foraging strategies of king penguins from Heard and Macquarie islands were compared using satellite telemetry, time-depth recorders and diet samples. Trip durations were 16.8±3.6 days and 14.8±4.1 days at Macquarie and Heard islands, respectively. At Macquarie Island, total distances travelled were 1281±203 km compared to 1425±516 km at Heard Island. The total time the penguins spent at sea was 393±66 h at Macquarie Island and 369±108 h at Heard Island. The penguins from Macquarie Island performed more deep dives than those from Heard Island. King penguins from Macquarie Island travelled 1.5±0.2 km h⁻¹ day⁻¹ compared to 1.3±0.1 km h⁻¹ day⁻¹. At Macquarie Island, 19% of dives were upto 70–90 m depth compared to 35% at Heard Island. The main dietary prey species were the fish Krefftychthis anderssoni and the squid Moroteuthis ingens in both groups. The differences in the at-sea distribution and the foraging behaviour of the two groups of penguins were possibly related to differences in oceanography and bathymetric conditions around the two islands. Dietary differences may be due to interannual variability in prey availability since the two colonies were studied during incubation but in different years.     

The metabolic cost of subcutaneous and abdominal rewarming in king penguins after long-term immersion in cold water

Marine endotherms in the polar regions face a formidable thermal challenge when swimming in cold water. Hence, they use morphological (fat, blubber) adjustment and peripheral vasoconstriction to reduce demands for heat production in water. The animals then regain normothermia when resting ashore. In the king penguin (Aptenodytes patagonicus) metabolic rate is lower in fed than in fasted individuals during subsequent rewarming on land. This has been suggested to be a consequence of diversion of blood flow to the splanchnic region in fed birds, which reduces peripheral temperatures. However, peripheral temperatures during recovery have never been investigated in birds with different nutritional status. The aim of this study was, therefore, to measure subcutaneous and abdominal temperatures during the rewarming phase on land in fasted and fed king penguins, and investigate to which extent any different rewarming were reflected in recovery metabolic rate (MR_R) after long term immersion in cold water. We hypothesized that fed individuals would have a slower increase of subcutaneous temperatures compared to fasted penguins, and a correspondingly lower MR_R. Subcutaneous tissues reached normothermia after 24.15 (back) and 21.36 min (flank), which was twice as fast as in the abdomen (46.82 min). However, recovery time was not affected by nutritional condition. MR_R during global rewarming (4.56 ± 0.42 W kg⁻¹) was twice as high as resting metabolic rate (RMR; 2.16 ± 0.59 W kg⁻¹). However, MR_R was not dependent on feeding status and was significantly elevated above RMR only until subcutaneous temperature had recovered. Contrary to our prediction, fed individuals did not reduce the subcutaneous circulation compared to fasted penguins and did not show any changes in MR_R during subsequent recovery. It seems likely that lower metabolic rate in fed king penguins on land reported in other studies might not have been caused primarily by increased circulation to the visceral organs.     

Diving patterns of breeding female rockhopper penguins (<Emphasis Type="Italic">Eudyptes chrysocome)</Emphasis>: Noir Island,Chile

The diving behaviour of female southern rockhopper penguins (Eudyptes chrysocome) was studied at Noir Island (54°30′S–73°00′W), Chile, in the southeast Pacific Ocean. This isolated island is located at the edge of the continental shelf in an area where the Humboldt Current originates, and holds a population of more than 150,000 breeding pairs. On 13 December 2005, four TDRs were successfully attached to females at the end of the brooding period and recorded diving activity at intervals of 2 s over the next 4 weeks. In total, 40 complete foraging trips were recorded. Trip duration was on average 42.4 ± 40.1 h and the proportion of overnight trips (60%) was the highest value found so far for this species. Mean dive depth and dive duration was 20.6 ± 19.4 m and 63.7 ± 36.4 s, respectively. The diving effort was higher than that of brooding females from the Indian Ocean and comparable with that of conspecifics from colonies in the southwest Atlantic in terms of diving rate (38 ± 14.2 dives h⁻¹), but slightly lower as regards the proportion of time spent underwater (61 ± 10.5%). This study confirms that the diving behaviour of rockhopper penguins varies as a function of the physical and biological characteristics of the foraging areas and of the particular stage of the breeding season.     

Foraging movements of Magellanic Penguins Spheniscus magellanicus in the Beagle Channel, Argentina, related to tide and tidal currents

This study investigates the movements of Magellanic Penguins Spheniscus magellanicus breeding on Isla Martillo during the early chick-rearing period. Foraging paths were reconstructed using GPS loggers that registered the penguins′ geographic position, water temperature and depth at regular intervals. The relationship between penguins′ movements and search strategies, tide and tidal currents were assessed. Mean trip duration was on average 14.7 ± 6.9 h (33% overnight), and the maximum distance reached was 24 ± 10 km. All penguins studied foraged to the east of the colony. We identified three phases based on the sinuosity and speed of the trajectory: transit, central and return. Foraging effort was higher during the central phase, followed by the transit phase, and lower in the return phase. Foraging success, measured as the percentage of time at the bottom during each phase, was also highest during the central phase. In all birds studied, the central phase of the foraging trip took place during ebb tide, and birds travelled to the foraging areas with flow tide running in the same direction of displacement. Our study suggests that penguins take advantage of tidal currents to facilitate their movements to and from the main foraging area, thereby reducing the energy expended. Moreover, we suggest that piscivorous diving birds may enhance their catch rate during ebb tide when fish are more concentrated near the channel bed.     

The Apache Longbow–Hellfire Missile Test at Yuma Proving Ground: Ecological Risk Assessment for Helicopter Overflight

A multi-stressor risk assessment was conducted at Yuma Proving Ground, Arizona, as a demonstration of the Military Ecological Risk Assessment Framework. The focus of the assessment was a testing program at Cibola Range, which involved an Apache Longbow helicopter firing Hellfire missiles at moving targets, that is, M60-A1 tanks. This article focuses on the wildlife risk assessment for the helicopter overflight. The primary stressors were sound and the view of the aircraft. Exposure to desert mule deer (Odocoileus hemionus crooki) was quantified using Air Force sound contour programs NOISEMAP and MR_NMAP, which gave very different results. Slant distance from helicopters to deer was also used as a measure of exposure that integrated risk from sound and view of the aircraft. Exposure-response models for the characterization of effects consisted of behavioral thresholds in sound exposure level or maximum sound level units or slant distance. Available sound thresholds were limited for desert mule deer, but a distribution of slant-distance thresholds was available for ungulates. The risk characterization used a weight-of-evidence approach and concluded that risk to mule deer behavior from the Apache overflight is uncertain, but that no risk to mule deer abundance and reproduction is expected.     

Massive breeding failures following a tsunami in a colonial seabird

Natural catastrophic events such as tsunamis may induce drastic decreases in breeding success of animal populations. We evaluate the impacts of flooding on the reproductive success of king penguins (Aptenodytes patagonicus) in the Crozet Archipelago. On 26 December 2004, a magnitude-9 earthquake created a large tsunami that flooded a colony at 6,500 km from the epicentre of the earthquake. On 30 January 2005, severe waves again flooded the colony. About 17–20% of the surface of the colony was impacted during each flood and 44% of the breeding birds abandoned their egg or chick following the two floodings. Although about 11% of birds laid another egg after the tsunami, none reproduced again after the second flood that happened later in the breeding season. Our results show that the tsunami directly affected the reproductive success of seabirds nesting near the coast.