How area restricted search of a pelagic seabird changes while performing a dual foraging strategy

Movements and foraging strategies of marine predators should cope with the hierarchical spatial distribution of resources. Therefore, in order to predict the at‐sea distribution of aerial predators, it is crucial to understand the factors governing trajectory decisions at different scales. Using first passage time (FPT) analysis on precision tracking information (GPS‐loggers data) we were able to examine the foraging strategy of Cory's shearwaters Calonectris diomedea and to detect the adoption of area‐restricted search (ARS), measuring the scale and duration of this behaviour. Data were collected from three different populations foraging in different oceanographic conditions. During long excursions birds only commuted between their colony and prey patches, while on their short movements birds increased the amount of looping movements. On short trips, birds addopted ARS behaviour at an average scale of 18 km and at a second nested scale of around 2 km. When engaging in long trips, first scale of ARS occurred on average at about 67 km of radii and than a second nested scale at a radii of 24 km. Overall, the different populations showed foraging patterns matching the habitats exploited: a) at smaller scales of ARS, sea‐surface temperature, chlorophyll‐a concentration and depth influenced the time of residence (i.e. FPT) of birds (with variations at a population level); b) at larger scales of ARS, FPT increased within regions of higher gradients of sea‐surface temperature, chlorophyl‐a concentration and depth. This study demonstrates that Cory's shearwaters adopt scale‐dependent adjustments of movement in relation to the hierarchical distribution of the environment they exploit, matching the scale and duration of ARS with the hierarchical distribution of the environmental features.     

Individual foraging site fidelity in lactating New Zealand fur seals: Continental shelf vs. oceanic habitats

Wide‐ranging marine central place foragers often exhibit foraging site fidelity to oceanographic features over differing spatial scales (i.e., localized coastal upwellings and oceanic fronts). Few studies have tested how the degree of site fidelity to foraging areas varies in relation to the type of ocean features used. In order to determine how foraging site fidelity varied between continental shelf and oceanic foraging habitats, 31 lactating New Zealand fur seals (Arctocephalus australis forsteri 1 ) were satellite tracked over consecutive foraging trips (14–108 d). Thirty‐seven foraging trips were recorded from 11 females that foraged on the continental shelf, in a region associated with a coastal upwelling, while 65 foraging trips were recorded from 20 females that foraged in oceanic waters. There were no significant differences in the mean bearings (to maximum distance) of individual's consecutive foraging trips, suggesting individual fidelity to foraging areas. However, overlap in area and time spent in area varied considerably between continental shelf and oceanic foragers. Females that foraged on the continental shelf had significantly greater overlap in consecutive foraging trips when compared to females that foraged in oceanic waters (overlap in 5 × 5 km grid cells visited on consecutive trips 55.9%± 20.4% and 13.4%± 7.6%, respectively). Females that foraged on the continental shelf also spent significantly more time within the same grid cell than females that foraged in oceanic waters (maximum time spent in 5 × 5 km grid cells: 14%± 5% and 4%± 2%, respectively). This comparatively high foraging site fidelity may reflect the concentration of productivity associated with a coastal upwelling system, the Bonney Upwelling. Lower foraging site fidelity recorded by seals that foraged in oceanic waters implies a lower density/larger scale habitat, where prey are more dispersed or less predictable at fine scales, when compared to the continental shelf region.     

Foraging movements of Leach's storm‐petrels Oceanodroma leucorhoa during incubation

Knowledge of foraging movements during the breeding season is key to understanding energetic stresses faced by seabirds. Using archival light loggers (geolocators), a Bayesian state–space model, and stable isotope analysis, we compared foraging movements of Leach's storm‐petrels Oceanodroma leucorhoa during their incubation periods in 2012 and 2013. Data were collected from two colonies, Bon Portage Island and Country Island, which are 380 km apart along the coast of Nova Scotia, Canada. Based on allometry for procellariiform mass, predicted foraging ranges for Leach's storm‐petrels are 200 km; however, observed maximum distances from the colony were 3 to 5 times that. Storm‐petrels from Country Island travelled 1015 ± 238 km southeast to the Laurentian fan and south of the Grand Banks whereas storm‐petrels from Bon Portage Island travelled 613 ± 167 km southeast, beyond the continental slope, east of Georges Bank. The average distance travelled in a return trip was 2287 ± 603 km and 1303 ± 351 km for Country Island and Bon Portage Island, respectively. There were no differences between years in cumulative distances travelled within islands, but foraging trips did not last as long in 2013 (4.7 ± 1.5 d) as they did in 2012 (6.2 ± 2.1 d). Stable isotope analyses indicated that, during the incubation period, prey items from Country Island were from higher trophic levels and possibly had higher energy content than those from Bon Portage Island, perhaps explaining the more distant and longer foraging trips for Country Island birds.     

Divergent diving behavior during short and long trips of a bimodal forager,the little auk Alle alle

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick‐ and self‐feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U‐shaped dives with slower ascent rates and shorter resting times (post‐dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near‐colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.     

Is food availability limiting African Penguins Spheniscus demersus at Boulders? A comparison of foraging effort at mainland and island colonies

The African Penguin Spheniscus demersus (Vulnerable) formed three new colonies during the 1980s, two on the South African mainland (Stony Point and Boulders) and one on Robben Island. One of the mainland colonies, at Boulders, Simon's Town, is in a suburban area, resulting in conflict with humans. Growth of the Boulders colony was initially rapid, largely through immigration, but has since slowed, possibly as a result of density‐dependent effects either on land (where there has been active management to limit the spread of the colony) or at sea. We test the latter hypothesis by comparing the foraging effort of Penguins feeding small chicks at island and mainland sites, and relate this to the foraging area available to birds. Three‐dimensional foraging paths of African Penguins were reconstructed using GPS and time–depth loggers. There were no intercolony differences in the rate at which birds dived during the day (33 dives/h), in diving depths (mean 17 m, max. 69 m) or in travelling speeds. The maximum speed recorded was 2.85 m/s, with birds travelling faster when commuting (average 1.18 m/s) than when foraging (0.93 m/s) or resting at sea (0.66 m/s during the day, 0.41 m/s at night). There were strong correlations between foraging trip duration, foraging range and total distance travelled. Foraging effort was correlated with chick age at Robben Island, but not at Boulders. Contrary to Ashmole's hypothesis, birds from Boulders (c. 1000 pairs) travelled further (46–53 km) and foraged for longer (13.2 h) than did birds from Robben Island (c. 7000 pairs) and Dassen Island (c. 21 000 pairs) (33 km, 10.3 h). The mean foraging range also differed significantly between mainland (18–20 km) and island colonies (9 km). The area available to central‐place‐foraging seabirds breeding on the mainland is typically less than that for seabirds breeding on islands, but the greater foraging range of Boulders birds results in an absolute foraging area roughly twice that of island colonies, and the area per pair is an order of magnitude greater for the relatively small Boulders colony. Ashmole's hypothesis assumes relatively uniform prey availability among colonies, but our results suggest this does not apply in this case. The greater foraging effort of Boulders birds probably reflects reduced prey availability in False Bay, and thus the recent slowing in growth at the colony may be the result of differential immigration rather than management actions to limit the spatial growth of the colony.     

Foraging strategy of the little auk Alle alle throughout breeding season – switch from unimodal to bimodal pattern

Energy and time allocation differs between incubation and chick‐rearing periods, which may lead to an adjustment in the foraging behaviour of parent birds. Here, we investigated the foraging behaviour of a small alcid, the little auk Alle alle during incubation and compared it with the chick‐rearing period in West Spitsbergen, using the miniature GPS (in Hornsund) and temperature loggers (in Magdalenefjorden). GPS‐tracking of 11 individuals revealed that during incubation little auks foraged 8–55 (median 46) km from the colony covering 19–239 (median 120) km during one foraging trip. Distance from the colony to foraging areas was similar during incubation and chick‐rearing period. During incubation 89% of foraging positions were located in the zone over shallower parts of the shelf (isobaths up to 200–300 m) with sea surface temperature below 2.5°C. Those environmental conditions are preferred by Arctic zooplankton community. Thus, little auks in the Hornsund area restrict their foraging (both during the incubation and chick‐rearing period) to the area under influence of cold, Arctic‐origin water masses where its most preferred prey, copepod Calanus glacialis is most abundant. The temperature logger data (from 4 individuals) indicate that in contrast to the chick‐rearing period, when parent birds alternated short and long trips, during the incubation they performed only long trips. Adopting such a flexible foraging strategy allows little auks to alter their foraging strategy to meet different energy and time demands during the two main stages of the breeding.     

Overnight foraging trips by chick‐rearing Nazca Boobies Sula granti and the risk of attack by predatory fish

Most tropical booby species complete breeding foraging trips within daylight hours, thus avoiding nights at sea. Nazca Boobies Sula granti are unusual in this respect, frequently spending one or more nights away from the nest. We used GPS dataloggers, time‐depth recorders, and changes in body weight to characterize foraging trips and to evaluate potential influences on the decisions of 64 adult Nazca Boobies to spend a night at sea, or to return to their chicks on Isla Española, Galápagos, in daylight hours. The tagged birds foraged east of Isla Española, undertaking both single‐day (2–15 h, 67% of trips) and overnight trips (28 h–7.2 days, 33%), and executing 1–19 foraging plunge‐dives per single‐day trip. Birds might forage longer if they are in nutritional stress when they depart, but body weight at departure was not correlated with trip length. Birds might be expected to return from longer trips with more prey for young, but they returned from single‐day and overnight trips with similar body weights, consistent with previous indications that Nazca Boobies forage until accumulating a target value of prey weight. Birds with a lower dive frequency during the first 5 h of a trip were more likely to spend the night at sea, suggesting that they might choose to spend the night at sea if prey capture success was low. At night, birds almost never dived and spent most of their time resting on the water’s surface (11.8–12.1 h, > 99% of the time between civil sunset and civil dawn). Thus, the night is an unproductive time spent among subsurface predators under low illumination. The birds’ webbed feet provided evidence of this risk: 24% of birds were missing > 25% of their foot tissue, probably due to attacks by predatory fish, and the amount of foot tissue lost increased with age, consistent with a cumulative risk across the lifespan. In contrast, other tropical boobies (Blue‐footed Sula nebouxii and Brown Boobies Sula leucogaster), which do not spend the night on the water, showed no such damage. These results suggest that chick‐rearing Nazca Boobies accept nocturnal predation risk on occasions of low prey encounter during a foraging trip’s first day.     

Disentangling olfactory and visual information used by field foraging birds

Foraging strategies of birds can influence trophic plant–insect networks with impacts on primary plant production. Recent experiments show that some forest insectivorous birds can use herbivore‐induced plant volatiles (HIPVs) to locate herbivore‐infested trees, but it is unclear how birds combine or prioritize visual and olfactory information when making foraging decisions. Here, we investigated attraction of ground‐foraging birds to HIPVs and visible prey in short vegetation on farmland in a series of foraging choice experiments. Birds showed an initial preference for HIPVs when visual information was the same for all choice options (i.e., one experimental setup had all options with visible prey, another setup with hidden prey). However, if the alternatives within an experimental setup included visible prey (without HIPV) in competition with HIPV‐only, then birds preferred the visual option over HIPVs. Our results show that olfactory cues can play an important role in birds’ foraging choices when visual information contains little variation; however, visual cues are preferred when variation is present. This suggests certain aspects of bird foraging decisions in agricultural habitats are mediated by olfactory interaction mechanisms between birds and plants. We also found that birds from variety of dietary food guilds were attracted to HIPVs; hence, the ability of birds to use plant cues is probably more general than previously thought, and may influence the biological pest control potential of birds on farmland.     

Time and travelling costs during chick‐rearing in relation to habitat quality in Little Owls Athene noctua

In many bird species, parents adjust their home‐ranges during chick‐rearing to the availability and distribution of food resources, balancing the benefits of energy intake against the costs of travelling. Over recent decades, European agricultural landscapes have changed radically, resulting in the degradation of habitats and reductions in food resources for farmland birds. Lower foraging success and longer foraging trip distances that result from these changes are often assumed to reduce the reproductive performance of parents, although the mechanisms are not well understood. We tested the behavioural response of chick‐rearing Little Owls Athene noctua to variation in habitat diversity in an agricultural landscape. We equipped females with GPS loggers and received adequate range‐use data for 19 individuals (6063–14 439 locations per bird). In habitats dominated by homogeneous cropland habitats, home‐ranges were over 12 ha in size, whereas in highly diverse habitats they were below 2 ha. Large home‐ranges were associated with increased flight activity (117% of that of birds in small home‐ranges) and distances travelled per night (152%), increased duration of foraging trips (169%) covering larger distances (246%), and reduced nest visiting rates (81%). The study therefore provides strong correlative evidence that Little Owls breeding in monotonous farmland habitats expend more time and energy for a lower benefit in terms of feeding rates than do birds in more heterogeneous landscapes. As nestling food supply is the main determinant of chick survival, these results suggest a strong impact of farmland characteristics on local demographic rates. We suggest that preserving and creating islands of high habitat diversity within uniform open agricultural landscapes should be a key target in the conservation of Little Owl populations.     

Visual tracking from a rigid‐hulled inflatable boat to determine foraging movements of breeding terns

ABSTRACT Defining the at‐sea foraging movements of seabirds is fundamental to understanding their ecology and can also be important in assessing the potential impact of marine developments such as offshore wind farms (OWFs). Surveys of predefined areas using aerial or boat‐based transect surveys may not allow adequate assessment of the relative importance of different areas to birds. Individual‐based satellite or radio‐telemetry can be effective in identifying foraging ranges and preferred areas, but may not be suitable for some species. We developed a method to determine the foraging movements of breeding terns (Sterna spp.) by visually tracking individuals using a rigid‐hulled inflatable boat (RHIB). Sandwich Terns (S. sandvicensis), Common Terns (S. hirundo), and Arctic Terns (S. paradisaea) were tracked from colonies in Norfolk and Anglesey, United Kingdom, from 2006 to 2008. The proportion of complete (from and to colony) trips varied from 29–60% among species, years, and colonies. Individual Sandwich Terns were tracked for periods up to 126 min over distances up to 72 km and as far as 54 km from the breeding colony, further than Arctic (up to 57 km and 29 km from the colony) and Common (to 29 km and <9 km from the colony) terns. Mean values were much smaller. Multivariate modeling of Sandwich Tern foraging trips indicated that flight speeds >50 km/hr coupled with greater distances from shore (>25 km) significantly reduced the likelihood of tracking a bird for an entire foraging trip. Use of different boats that differ in speed and performance may alleviate such issues. Visual tracking allowed us to collect data on foraging behavior, flight height, and prey capture rates, and also permitted comparisons between species. Our results indicate that visual tracking may be an effective means of determining the foraging movements and at‐sea behavior of a variety of short‐ranging, day‐active seabirds.     

Foraging by little auks in the distant marginal sea ice zone during the chick-rearing period

During the chick-rearing period, little auks Alle alle adopt a bimodal foraging strategy, alternating long trips with several short ones. It has been postulated that they reach more remote areas during long feeding trips than during short ones. However, the range of their foraging flights has never actually been measured. The aims of this study were to find the exact location of the little auk feeding grounds and to investigate whether they reach remote areas during long foraging trips using miniature GPS and temperature loggers. The study was conducted in 2009 in Magdalenefjorden (79°34′N, 11°04′E), one of the main breeding grounds of little auks on Spitsbergen. The temperature logger records indicated that during short trips, little auks visit warmer waters (situated close to the colony) than during long ones. The tracks of two GPS-equipped birds indicated that during long trips little auks foraged in the distant, food-abundant marginal sea ice zone, at least 100 km away from the colony. During long trips, birds make several stops at sea, perhaps sampling the foraging area with respect to prey distribution. Since food conditions near the studied colony are usually suboptimal, little auks may be exploiting distant feeding areas to compensate for the poorer-quality food available at nearby foraging grounds. The extended duration of long foraging trips may enable birds to collect food for chicks on food-abundant, remote foraging grounds as well as acquire, process and excrete food needed for self-maintenance, reducing the costs of flight to the colony.     

Does prey capture induce area-restricted search? A fine-scale study using GPS in a marine predator, the wandering albatross

In a patchy environment, predators are expected to increase turning rate and start an area-restricted search (ARS) when prey have been encountered, but few empirical data exist for large predators. By using GPS loggers with devices measuring prey capture, we studied how a marine predator adjusts foraging movements at various scales in relation to prey capture. Wandering albatrosses use two tactics, sit and wait and foraging in flight, the former tactic being three times less efficient than the latter. During flight foraging, birds caught large isolated prey and used ARS at scales varying from 5 to 90 km, with large-scale ARS being used only by young animals. Birds did not show strong responses to prey capture at a large scale, few ARS events occurred after prey capture, and birds did not have high rates of prey capture in ARS. Only at small scales did birds increase sinuosity after prey captures for a limited time period, and this occurred only after they had caught a large prey item within an ARS zone. When this species searches over a large scale, the most effective search rule was to follow a nearly straight path. ARS may be used to restrict search to a particular environment where prey capture is more predictable and profitable.     

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.     

Hierarchical patch dynamics and animal movement pattern

In hierarchical patch systems, small-scale patches of high density are nested within large-scale patches of low density. The organization of multiple-scale hierarchical systems makes non-random strategies for dispersal and movement particularly important. Here, we apply a new method based on first-passage time on the pathway of a foraging seabird, the Antarctic petrel (Thalassoica antarctica), to quantify its foraging pattern and the spatial dynamics of its foraging areas. Our results suggest that Antarctic petrels used a nested search strategy to track a highly dynamic hierarchical patch system where small-scale patches were congregated within patches at larger scales. The birds searched for large-scale patches by traveling fast and over long distances. Once within a large-scale patch, the birds concentrated their search to find smaller scale patches. By comparing the pathway of different birds we were able to quantify the spatial scale and turnover of their foraging areas. On the largest scale we found foraging areas with a characteristic scale of about 400 km. Nested within these areas we found foraging areas with a characteristic scale of about 100 km. The large-scale areas disappeared or moved within a time frame of weeks while the nested small-scale areas disappeared or moved within days. Antarctic krill (Euphausia superba) is the dominant food item of Antarctic petrels and we suggest that our findings reflect the spatial dynamics of krill in the area.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Foraging movements of emperor penguins at Pointe Géologie, Antarctica

The foraging distributions of 20 breeding emperor penguins were investigated at Pointe Géologie, Terre Adélie, Antarctica by using satellite telemetry in 2005 and 2006 during early and late winter, as well as during late spring and summer, corresponding to incubation, early chick-brooding, late chick-rearing and the adult pre-moult period, respectively. Dive depth records of three post-egg-laying females, two post-incubating males and four late chick-rearing adults were examined, as well as the horizontal space use by these birds. Foraging ranges of chick-provisioning penguins extended over the Antarctic shelf and were constricted by winter pack-ice. During spring ice break-up, the foraging ranges rarely exceeded the shelf slope, although seawater access was apparently almost unlimited. Winter females appeared constrained in their access to open water but used fissures in the sea ice and expanded their prey search effort by expanding the horizontal search component underwater. Birds in spring however, showed higher area-restricted-search than did birds in winter. Despite different seasonal foraging strategies, chick-rearing penguins exploited similar areas as indicated by both a high ‘Area-Restricted-Search Index’ and high ‘Catch Per Unit Effort’. During pre-moult trips, emperor penguins ranged much farther offshore than breeding birds, which argues for particularly profitable oceanic feeding areas which can be exploited when the time constraints imposed by having to return to a central place to provision the chick no longer apply.     

At-sea distribution and scale-dependent foraging behaviour of petrels and albatrosses: a comparative study

1. In order to study and predict population distribution, it is crucial to identify and understand factors affecting individual movement decisions at different scales. Movements of foraging animals should be adjusted to the hierarchical spatial distribution of resources in the environment and this scale-dependent response to environmental heterogeneity should differ according to the forager's characteristics and exploited habitats. 2. Using First-Passage Time analysis, we studied scales of search effort and habitat used by individuals of seven sympatric Indian Ocean Procellariiform species fitted with satellite transmitters. We characterized their search effort distribution and examined whether species differ in scale-dependent adjustments of their movements according to the marine environment exploited. 3. All species and almost all individuals (91% of 122 individuals) exhibited an Area-Restricted Search (ARS) during foraging. At a regional scale (1000s km), foraging ranges showed a large spatial overlap between species. At a smaller scale (100s km, at which an increase in search effort occurred), a segregation in environmental characteristics of ARS zones (where search effort is high) was found between species. 4. Spatial scales at which individuals increased their search effort differed between species and also between exploited habitats, indicating a similar movement adjustment for predators foraging in the same habitat. ARS zones of the two populations of wandering albatross Diomedea exulans (Crozet and Kerguelen) were similar in their adjustments (i.e. same ARS scale) as well as in their environmental characteristics. These two populations showed a weak spatial overlap in their foraging distribution, with males foraging in more southerly waters than females in both populations. 5. This study demonstrates that predators of several species adjust their foraging behaviour to the heterogeneous environment and these scale-dependent movement adjustments depend on both forager and environment characteristics.     

Threshold foraging by gray whales in response to fine scale variations in mysid density

The gray whale (Eschrichtius robustus) is a coastal species whose nearshore summer foraging grounds off the coast of British Columbia offer an opportunity to study the fine scale foraging response of baleen whales. We explore the relationship between prey density and gray whale foraging starting with regional scale (10 km) assessments of whale density (per square kilometer) and foraging effort as a response to regional mysid density (per cubic meter), between 2006 and 2007. In addition we measure prey density at a local scale (100 m), while following foraging whales during focal surveys. We found regional mysid density had a significant positive relationship with both gray whale density and foraging effort. We identify a threshold response to regional mysid density for both whale density and foraging effort. In 2008 the lowest average local prey density measured beside a foraging whale was 2,300 mysids/m³. This level was maintained even when regional prey density was found to be substantially lower. Similar to other baleen whales, the foraging behavior of gray whales suggests a threshold response to prey density and a complex appreciation of prey availability across fine scales.     

Quantitative analysis of bottlenose dolphin movement patterns and their relationship with foraging

1. Broad-scale telemetry studies have greatly improved our understanding of the ranging patterns and habitat-use of many large vertebrates. However, there often remains considerable uncertainty over the function of different areas or the factors influencing habitat selection. Further insights into these processes can be obtained through analyses of finer scale movement patterns. For example, search behaviour may be modified in response to prey distribution and abundance. 2. In this study, quantitative analysis techniques are applied to the movements of bottlenose dolphins, recorded from land using a theodolite, to increase our understanding of their foraging strategies. Movements were modelled as a correlated random walk (CRW) and a biased random walk (BRW) to identify movement types and using a first-passage time (FPT) approach, which quantifies the time allocated to different areas and identifies the location and spatial scale of intensive search effort. 3. Only a quarter of the tracks were classed as CRW movement. Turning angle and directionality appeared to be key factors in determining the type of movement adopted. A high degree of overlap in search effort between separate movement paths indicated that there were small key sites (0.3 km radius) within the study area (4 km(2)). Foraging behaviour occurred mainly within these intensive search areas, indicating that they were feeding sites. 4. This approach provides a quantitative method of identifying important foraging areas and their spatial scale. Such techniques could be applied to movement paths for a variety of species derived from telemetry studies and increase our understanding of their foraging strategies.     

Inter-population variation in the behaviour of adult and juvenile Red-footed Boobies Sula sula

Early life is a critical phase of the life cycle of animals and is attracting increased attention because little information is available on the behaviour of young individuals during this period. Behaviour during early life is probably influenced by the environmental conditions encountered by young animals, but data on intraspecific variation between breeding sites during this crucial period of life are limited. Here we study variability in the foraging behaviour of juveniles and adults in three colonies of a pantropical seabird, the Red-footed Booby Sula sula. Both adults and juveniles were measured and fitted with GPS loggers in three remote islands: Genovesa (Galapagos, Eastern Pacific Ocean), Europa (Western Indian Ocean) and Surprise (New Caledonia, Western Pacific Ocean). Foraging behaviour was compared between age-classes, sex and colonies by examining trip characteristics, different behaviours at sea, potential associations between individuals and morphological characteristics. Compared with adults, juveniles conducted shorter trips that were restricted to around the colony, especially on Genovesa (max. range: 203.4 ± 125.1 km and 3.6 ± 3.1 km, respectively). Juveniles appeared more constrained by poor flight skills and experience rather than by their morphology. Adults travelled 45% of the time during at-sea trips, whereas juveniles spent a a lower proportion of time travelling but foraged more often using an ‘area-restricted search’ behaviour, potentially training to catch prey. Associations between juveniles were commonly detected in the three colonies and occurred mostly during foraging, suggesting that social learning is an important strategy. Variability of morphometric measurements in both adults and juveniles was high between sites, with larger birds found on Genovesa. These results suggest that adaptations to local environmental conditions are already visible in their early life. Future studies should continue to investigate the behavioural flexibility of juvenile birds to better understand the effect of local environmental conditions during this critical stage of life.     

Quantifying the effects of habitat structure on prey detectability and accessibility to farmland birds

For species that rely on visual cues to detect prey items, increasing the structural complexity of a patch can greatly influence forager behaviour through consequent reductions in prey detectability and accessibility. These effects are likely to manifest themselves in terms of foraging site selection and there is plentiful evidence for preferential site selection for a suite of taxa. However, the underlying effects of habitat structure on foraging behaviour, which are likely to drive these observed site selections, are much less well understood. We present the results of two studies designed to quantify the effects of vegetation structure on prey detectability and accessibility to avian invertebrate feeders and granivores on farmland. There was a significant negative relationship between potential prey detectability and both distance and vegetation height in cereal crops and stubbles for Northern Lapwings Vanellus vanellus . The interscan distance travelled by Lapwings differed significantly between habitats, with longer distances travelled in cereal crops and harrowed compared with ploughed soil and grasses. The peck rate, head-up rate and mean search period of foraging Chaffinches Fringilla coelebs were not affected by increasing vegetation structure but forager mobility was significantly reduced. We hope that by quantifying the effects of vegetation structure on prey detectability and accessibility we can highlight the importance of considering these factors, as well as prey abundance, when developing management strategies for farmland birds.