Hardly hard-hearted: heart rate responses of incubating Northern Giant Petrels (Macronectes halli) to human disturbance on sub-Antarctic Marion Island

Guidelines for visitors to sub-Antarctic Marion Island recommend 15 and 100 m minimum approach distances for breeding Northern (Macronectes halli) and Southern (Macronectes giganteus) Giant Petrels, respectively. Using artificial eggs containing FM transmitters, we measured the heart rate responses of incubating Northern Giant Petrels to pedestrian approaches. The mean resting heart rate was 80 beats per minute. Heart rates increased upon the detection of a person approximately 40 m away, and continued to increase during the approach to 5 m. Maximum increases over resting heart rate in response to natural disturbances and human approach were 97 and 204%, respectively. Northern Giant Petrels appear at least as sensitive to human disturbance as their congenerics. While low-key disturbance is unlikely to affect this solitary breeder as severely as it would the colonial Southern Giant Petrel, improved protection from disturbance could be achieved by restricting human passage through breeding colonies of Northern Giant Petrels to defined paths.     

Female and male Leach's Storm Petrels (Hydrobates leucorhous) pursue different foraging strategies during the incubation period

Reproduction in procellariiform birds is characterized by a single egg clutch, slow development, a long breeding season and obligate biparental care. Female Leach's Storm Petrels Hydrobates leucorhous, nearly monomorphic members of this order, produce eggs that are between 20 and 25% of adult bodyweight. We tested whether female foraging behaviour differs from male foraging behaviour during the ~ 44-day incubation period across seven breeding colonies in the Northwest Atlantic. Over six breeding seasons, we used a combination of Global Positioning System and Global Location Sensor devices to measure characteristics of individual foraging trips during the incubation period. Females travelled significantly greater distances and went farther from the breeding colony than did males on individual foraging trips. For both sexes, the longer the foraging trip, the greater the distance. Independent of trip duration, females travelled farther, and spent a greater proportion of their foraging trips prospecting widely, as defined by behavioural categories derived from a hidden Markov Model. For both sexes, trip duration decreased with date. Sex differences in these foraging metrics were apparently not a consequence of morphological differences or spatial segregation. Our data are consistent with the idea that female foraging strategies differed from male foraging strategies during incubation in ways that would be expected if females were still compensating for egg formation.     

Sexual dimorphism and sexual segregation in foraging strategies of northern giant petrels, Macronectes halli, during incubation

Giant petrels ( Macronectes spp.) are the most sexually dimorphic of all seabirds. We used satellite-tracking and mass change during incubation to investigate the influence of sexual size dimorphism, in terms of the intersexual food competition hypothesis, on foraging and fasting strategies of northern giant petrels at South Georgia. Females foraged at sea whereas males foraged mainly on the South Georgia coast, scavenging on seal and penguin carcasses. Foraging effort (flight speed, distance covered, duration of foraging trips) was greater for females than for males. In contrast, foraging efficiency (proportionate daily mass gain while foraging) was significantly greater for males than for females. Females were significantly closer to the desertion mass threshold than males and could not compensate for the mass loss during the incubation fast while foraging, suggesting greater incubation costs for females than for males. Both sexes regulated the duration and food intake of foraging trips depending on the depletion of the body reserves. In males the total mass gain was best explained by mass at departure and body size. We suggest that sexual segregation of foraging strategies arose from size-related dominance at carcasses, promoting sexual size dimorphism. Our results indicate that sex-specific differences in fasting endurance, contest competition over food and flight metabolic rates are key elements in maintenance of sexual size dimorphism, segregating foraging strategies and presumably reducing competition between sexes.     

Foraging behaviour and habitat partitioning of two sympatric cormorants in Patagonia, Argentina

Radiotelemetry was used to assess the distribution and diving behaviour of Rock Shags Phalacrocorax magellanicus and Red-legged Cormorants Phalacrocorax gaimardi breeding in sympatry, and Rock Shags breeding in isolation. When breeding in sympatry there was little overlap in the foraging locations of the two species, with the highest densities of each species separated by 10 km. Red-legged Cormorants fed significantly closer to the breeding colony than did Rock Shags and undertook shorter foraging trips, making almost twice as many foraging trips per day as Rock Shags. Rock Shags breeding in isolation had a shorter foraging range than the birds breeding in sympatry with Red-legged Cormorants and foraging trip duration was significantly shorter. However, the number of feeding trips per day was similar between areas of sympatry and allopatry. Differences in the foraging ecology of Rock Shags in areas of sympatry and allopatry may be due to interspecific competition, which forces niche differentiation. The distance between foraging sites, the speed of movement of the prey, a species tendency to move into prey-depleted areas and the length of the breeding season (during which the birds are constrained to be in the same area) may play critical roles in determining the extent to which differential area use by competitors is a strategy that benefits both parties.     

High foraging site fidelity and spatial segregation among individual great black‐backed gulls

Individual foraging site fidelity, whereby individuals repeatedly visit the same foraging areas, is widespread in nature, and likely benefits individuals through higher foraging efficiency and potentially, higher breeding success. It may arise as a consequence of habitat or resource specialisation, or alternatively, where resources are abundant or predictable, the partitioning of space might guarantee individuals exclusive foraging opportunities. We tracked seven adult great black‐backed gulls Larus marinus at a North Sea colony from early incubation to the end of the breeding season in 2016, providing a total of 1170 foraging trips over a mean ± SD tracking period of 67 ± 16 days. There was clear spatial segregation between individuals, with almost no overlap of their core areas (50% utilisation distribution) during incubation and chick‐rearing. Core areas were relatively small and there was high repeatability (R ± SE) in foraging parameters, including initial departure direction (0.73 ± 0.11), foraging range (0.41 ± 0.14) and cumulative distance travelled (0.19 ± 0.1) throughout the breeding season. Despite the low spatial overlap, there was little evidence of differential habitat use by individuals. The near‐exclusive individual foraging areas of this species, usually considered to be a generalist, indicate that where there is high resource availability throughout the breeding season and a small local population, individuals appear to adopt a territorial strategy which likely reduces intraspecific competition.     

The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island

While ashore King Penguins Aptenodytes patagonicus are fed upon by a guild of five predator-scavenger seabirds. During the winter (April-October) male Southern Giant Petrels Macronectes giganteus killed an estimated 6430 (11.2%) of Marion Island's King Penguin chicks, although most birds only scavenged in the colonies. The rate of predation varied, with peaks in April-May and in September. The proportion of successful attacks was 22.7%. There was a strong correlation between colony size and the rate of accumulation of chick corpses. Kelp Gulls Larus dominicanus and Lesser Sheathbills Chionis minor also scavenged penguin corpses but Northern Giant Petrels M. halli and female Southern Giant Petrels rarely entered the colonies. During the summer predation was mainly by Sub-Antarctic Skuas Catharacta lonnbergi which took eggs and small chicks.     

Foraging location and range of White-chinned Petrels Procellaria aequinoctialis breeding in the South Atlantic

The foraging range and principal feeding areas of White‐chinned Petrels breeding at South Georgia were determined using satellite telemetry. Foraging trips during incubation lasted 12–15 days and covered 3000–8000 km and 2–11 days and 1100–5900 km during chick‐rearing. Adults covered less distance per day during chick‐rearing (71 km) than during incubation (91 km) but the proportion covered at night (47%) was the same. Mean (31–34 km/h) and maximum (80 km/h) flight velocities were similar during both periods of the breeding season and during day and night. Between incubation shifts, White‐chinned Petrels travelled to the Patagonian shelf; during chick‐rearing they foraged more extensively. Most locations were between 30° to 55°W and 52° to 60°W around South Georgia/Shag Rocks and south to the South Orkney Islands. Diet samples from known foraging locations suggested birds fed mainly on krill and squid. They caught the squid Brachioteuthis? picta and Galiteuthis glacialis around Shag Rocks/South Georgia and also at sites close to the South Orkney Islands; Illex argentinus on the Patagonian shelf. Dispersal of adults after breeding failure was south to the South Orkney Islands then west to the Falkland Islands. This study confirms that breeding White‐chinned Petrels are amongst the widest‐ranging of seabirds; they may minimise competition with other Procellariiformes in the South Atlantic by their more extensive foraging range. The nature and extent of their range also brings substantial risk of high mortality rate in South Atlantic long‐line fisheries.     

Satellite tracking of Bulwer's Petrels Bulweria bulwerii in the Canary Islands

Capsule?The breeding foraging and post-breeding dispersal movements of five satellite-tagged Bulwer's Petrels from the Canary Islands were recorded. Foraging trips lasted 5.6 days in average (n?=?3), while the mean distance covered was 1261 km, and foraging areas were located over the continental slope and the adjacent pelagic waters, around 1200–2000 m depth and up to 350 km from the colony. After the chick-rearing period, birds dispersed southwestwards to the tropical waters of the central Atlantic.     

A further contribution on the biology of the Antarctic flea,Glaciopsyllus antarcticus (Siphonaptera: Ceratophyllidae)

Summary Experiments showed that adult Antarctic Fleas Gladopsyllus antarcticus preferred dark over light conditions, dry over wet conditions, and fine substrate over coarse substrate. Examination of seabird chicks and collections of nest material indicated that the Southern Fulmar Fulmarus glacialoides, is the major host species of the Antarctic Flea, while Snow Petrels Pagodroma nivea, Cape Petrels Daption capense, Antarctic Petrels Thalassoica antarctica, and Wilson's Storm-Petrels Oceanites oceanicus are minor hosts. This is the first report of Antarctic Fleas occuring on either Antarctic Petrels or Wilson's Storm-Petrels. No fleas were found associated with Southern Giant Petrels Macronectes giganteus, Antarctic Skuas Catharacta maccormicki, or Adelie Penguins Pygoscelis adeliae. No live fleas were found in Southern Fulmar nest material during their period of dispersal over the winter months, supporting the hypothesis that Antarctic Fleas survive the winter period by remaining on their seabird host.     

Competition between perch (Perca fluviatilis) and ruffe (Gymnocephalus cernuus): the advantage of turning night into day

1. The outcome of interspecific competition for food resources depends both on the competitors’ sensory abilities and on environmental conditions. In laboratory experiments we tested the influence of daylight and darkness on feeding behaviour and specific growth rate (SGR) of two species with different sensory abilities. 2. We used perch (Perca fluviatilis) as a visually orientated, and ruffe (Gymnocephalus cernuus) as a mechano‐sensory oriented predator and tested their growth rates and behaviour under conditions of interspecific and intraspecific competition. Three different foraging conditions were used: food supplied (i) only during the day, (ii) only during the night or (iii) during both day and night. 3. In perch neither SGR nor feeding behaviour were influenced substantially by interspecific competition during daylight. During darkness their foraging behaviour changed markedly and their access to the food source as well as their SGR were negatively affected by the presence of ruffe. 4. Ruffe's foraging behaviour did not change during either day or night with interspecific competition. During the night ruffe's SGR was higher with interspecific competition, probably because of a release from intraspecific competition and the competitive inferiority of perch during the night. 5. Because of its seonsory abilities ruffe feeds predominantly at night, thereby reducing competitive interference from perch.     

Year‐round spatial movements and trophic ecology of Trindade Petrels (Pterodroma arminjoniana)

Trindade Petrels (Pterodroma arminjoniana) are vulnerable gadfly petrels that breed on the remote Trindade Island, located ~1100 km off the Brazilian coast. Little is known about their spatial ecology, and their trophic ecology has only been described for the breeding season. We tagged four Trindade Petrels with global location sensing loggers (GLS) from October 2013 to November 2014 and sampled the blood and feathers (innermost primary and the eighth secondary) of 14 individuals to evaluate their year‐round spatial and isotopic ecology. We examined individual distributions, habitat use and suitability, activity, and isotopic values during the breeding, migration, and non‐breeding periods. Trindade Petrels used areas in the southwest Atlantic Ocean (between 10°N and 50°S in latitude) during the breeding season. They migrated through pelagic waters of the tropical Atlantic to the northwest Atlantic, where they spent the non‐breeding season. Trindade Petrels used mostly tropical to subtropical waters in areas of intermediate to high wind speeds and low marine productivity. Individuals spent more time foraging at night than during the day. During the breeding season, birds in northerly areas had higher carbon‐13 values, and birds that used more pelagic areas foraged on prey at a higher trophic level (higher nitrogen‐15 values) than those in more southern and coastal areas. Isotopic values during the breeding, migration, and non‐breeding periods differed, possibly due to differences among individuals in their at‐sea distribution throughout the year. We confirmed the non‐breeding distribution of Trindade Petrels, which was previously known only from vessel sightings and stranded birds. Our results also suggest a strong temporal segregation in the at‐sea distribution and trophic ecology between two groups of individuals, which might indicate the existence of two separate breeding populations.     

Breeding strategies of Antarctic Petrels Thalassoica antarctica and Southern Fulmars Fulmarus glacialoides in the high Antarctic and implications for reproductive success

Breeding strategies of two closely related fulmarine petrels were studied on Ardery Island, on the continental coast of East Antarctica, where short summers are expected to narrow the time-window for reproduction. Both species had a similar breeding period (97 days from laying to fledging) but Antarctic Petrels Thalassoica antarctica bred up to 16 days earlier than Southern Fulmars. During the pre-laying exodus, all Antarctic Petrels deserted the colony, whereas some Southern Fulmars Fulmarus glacialoides remained. Antarctic Petrels exhibited stronger synchronization in breeding, made longer foraging trips and spent less time guarding their chicks than Southern Fulmars. Overall breeding success of both species was similar but failures of Antarctic Petrels were concentrated in the early egg-phase and after hatching, when parents ceased guarding. Southern Fulmars lost eggs and chicks later in the breeding cycle and so wasted more parental investment in failed breeding attempts. Different breeding strategies may be imposed by flight characteristics; Southern Fulmars are less capable of crossing large expanses of pack ice and need to delay breeding until the sea ice retreats and breaks up. However, due to the short summer they risk chick failure when weather conditions deteriorate late in the season.     

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.     

Specificity of grouping behaviour: comparing colony sizes for the same seabird species in distant populations

Animal collective patterns such as group size frequency distributions often show substantial intraspecific variation, suggesting low species‐specific consistency. Here, we dissect intraspecific vs interspecific components of colony size variation to estimate the repeatability (R) of colony size frequency distribution (CSFD) statistics for seabird species breeding in at least two out of four distant geographic areas of the Northern Hemisphere (21 species; 57 populations; 21 665 colonies; 9 326 479 breeding pairs). Colony sizes were highly variable both within and between species. We estimated the proportion of between‐species variation using the repeatability statistic. Colony size‐related statistics of CSFDs (e.g. geometric mean) showed high repeatabilities (R = 0.73–0.88), and shape‐related measures ranged from null (kurtosis), moderate (fit to a log‐normal distribution, R = 0.62) to highly repeatable (e.g. skewness, R = 0.74–0.87). We thus show that species collective patterns can be at the same time highly variable within species and a robust species‐specific trait that bridge ecological spatio‐temporal heterogeneities.     

Spatio-temporal overlap between the at-sea distribution of Southern Giant Petrels and fisheries at the Patagonian Shelf

The interactions between seabirds and fisheries pose significant threats for the seabird species such as incidental capture. In contrast, several species of seabirds meet part of their energetic requirements through the use of fisheries discards. Knowledge about the relationship between at-sea distribution of Procellariiformes and fisheries is a key tool in marine ecosystem management. We analysed the spatio-temporal relationship between the areas used by 16 satellite-tracked breeding adults of the Southern Giant Petrel and fisheries distribution and catch at the Patagonian Shelf. We also determined the time spent by adults in different marine jurisdictions. Results indicated a marked spatio-temporal association between birds and fisheries, mainly trawlers. The Southern Giant Petrels concentrated their foraging effort over Argentinean waters. The use of an abundant and predictable food source provided by the fisheries discards may be one of the factors affecting the dynamics of the Southern Giant Petrel populations in Patagonia, Argentina.     

Temperature modulates the effects of predation and competition on mosquito larvae

1. Predation risk affects interspecific competition by decreasing foraging activity and relative competitive ability. Predation risk is determined by predators' prey choice and prey responses, both of which can be influenced by temperature. Temperature is especially important for larval prey and can result in a trade‐off between predator‐induced decreases in foraging activity and growth. Interspecific competition must also be examined in relation to intraspecific density‐dependent competition; weaker interspecific competition leads to coexistence of competitors. 2. This study explored how temperature (15 and 25 °C) could affect a focal species, larvae of the mosquito Culex quinquefasciatus, by examining prey choice in a shared predator (mosquitofish; Gambusia holbrooki) and the effects of predation risk on interspecific competition with Limnodynastes peronii tadpoles. Intraspecific density‐dependent competition in C. quinquefasciatus at these temperatures was also examined. 3. At 25 °C, G. holbrooki consumption of both C. quinquefasciatus and L. peronii increased; however, the effects of interspecific competition on mosquito survival did not decrease with L. peronii exposure to predation risk. The relationship between intraspecific density‐dependent competition and interspecific competition was temperature‐dependent, with competitive dominance of L. peronii at 25 °C. Male and female mosquitoes had different temperature‐dependent responses, indicating sex‐specific intrinsic responses to starvation and differential selection pressures. At 25 °C, females were susceptible to interspecific competition by L. peronii, while males were susceptible to intraspecific competition. 4. The use of competitors as biological controls has implications for mosquito disease transmission, and these results suggest that control effectiveness may be modified by climate change.     

Strong temporal consistency in the individual foraging behaviour of Imperial Shags Phalacrocorax atriceps

Individual consistency in foraging behaviour can generate behavioural variability within populations and may, ultimately, lead to species diversification. However, individual‐based long‐term behavioural studies are particularly scarce in seabird species. Between 2008 and 2011, breeding Imperial Shags Phalacrocorax atriceps at the Punta León colony, Argentina, were tracked with GPS devices to evaluate behavioural consistency during their foraging trips. Within a breeding season, individuals were highly consistent in the maximum distances they reached from the shore and the colony, as well as in the time invested in flight and diving across consecutive days during early chick rearing. In addition, each individual had its specific foraging area distinct from the foraging area of other individuals. Comparing between early and late chick rearing in the same season, individuals were consistent, to a lesser degree, in the maximum distance they reached from the colony and the shore, increasing in consistency later on in the season. Within the season, females were more consistent than males in the maximum distance they moved from the colony and the shore, the sexes segregated in their foraging areas and individual females were segregated from one another. Twenty‐eight individuals tracked in different breeding seasons were marginally consistent in their trip durations and maximum distance reached from shore across seasons. Among seasons, foraging locations differed between sexes and among individual females. Individuals from this colony exhibited consistency over time in several aspects of foraging behaviour, which may be due to a combination of individual characteristics such as learning abilities, breeding experience or health, as well as targeted prey type and stability of the environment at this location.     

Tracking reveals behavioural coordination driven by environmental constraints in the Black-vented Shearwater Puffinus opisthomelas

During the breeding season, seabird foraging trips are constrained by nest attendance schedule and are necessarily colony centred. Oceanographic cues play a major role in the choice of foraging areas to minimize the time spent away from the nest. Here, we analysed the foraging tracks of Black-vented Shearwaters Puffinus opisthomelas during the incubation and chick-rearing periods of 2016 and 2017 at Isla Natividad (Mexico). We applied expectation-maximization binary clustering to track data to clusterize different behaviour patterns during foraging flights. We then applied binary generalized linear mixed models to characterize of foraging areas based on of environmental variables. We finally used kernel estimation techniques to describe main foraging areas. In 2016, breeding shearwaters used two core areas for foraging and resting on the water; the core area delineated by males was located northward from the colony in the Vizcaino Bay and the core area for females was located southward from the colony at the entrance of San Ignacio Lagoon. In 2017, males and females used the same areas with no evident segregation. Our study provided the first information on Black-vented Shearwater foraging areas during the breeding season and indicated that sexual segregation within coastal waters off the central Baja California Peninsula might be a foraging strategy during years of warmer ocean, likely less productive regimes. Factors including ocean-climate-mediated sexual segregation at sea, leading to interannual variation in foraging areas, should be considered when evaluating management actions intended to protect critical foraging habitats for Black-vented Shearwaters.     

Investigations of density interactions among breeding birds in ponderosa pine forests: correlative and experimental evidence

Summary We assessed the importance of interspecific competition among insectivorous birds breeding in northcentral Arizona's ponderosa pine forests. We examined density interactions among species using two analytic approaches; correlative and experimental. The correlative approach examined patterns of change in breeding densities over four years at the community level and within two foraging guilds; picker-gleaners and aerial feeders. The relationships between morphological and behavioral similarity with pairwise density interactions were also assessed. Our experimental approach involved placement of nest boxes on two treatment plots to increase breeding densities of secondary cavity nesting birds that were in foraging guilds with open nesting insectivores.We found little evidence of interspecific competition. Patterns of density fluctuations indicated large positive covariances among species at both the community level and within guilds. Pairwise density interactions were independent of morphological or behavioral similarity. Nest boxes significantly increased breeding densities of the secondary cavity nesters. However, these increases did not induce reprocal density changes in the open nesting species. Interspecific competition for food during the breeding season appears to be unimportant in ponderosa pine bird communities.     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.