David Grémillet

Climate change and the ecology and evolution of Arctic vertebrates

Climate change is taking place more rapidly and severely in the Arctic than anywhere on the globe, exposing Arctic vertebrates to a host of impacts. Changes in the cryosphere dominate the physical changes that already affect these animals, but increasing air temperatures, changes in precipitation, and ocean acidification will also affect Arctic ecosystems in the future. Adaptation via natural selection is problematic in such a rapidly changing environment. Adjustment via phenotypic plasticity is therefore likely to dominate Arctic vertebrate responses in the short term, and many such adjustments have already been documented. Changes in phenology and range will occur for most species but will only partly mitigate climate change impacts, which are particularly difficult to forecast due to the many interactions within and between trophic levels. Even though Arctic species richness is increasing via immigration from the South, many Arctic vertebrates are expected to become increasingly threatened during this century.

Marine no-take zone rapidly benefits endangered penguin

No-take zones may protect populations of targeted marine species and restore the integrity of marine ecosystems, but it is unclear whether they benefit top predators that rely on mobile pelagic fishes. In South Africa, foraging effort of breeding African penguins decreased by 30 per cent within three months of closing a 20 km zone to the competing purse-seine fisheries around their largest colony. After the fishing ban, most of the penguins from this island had shifted their feeding effort inside the closed area. Birds breeding at another colony situated 50 km away, whose fishing grounds remained open to fishing, increased their foraging effort during the same period. This demonstrates the immediate benefit of a relatively small no-take zone for a marine top predator relying on pelagic prey. Selecting such small protected areas may be an important first conservation step, minimizing stakeholder conflicts and easing compliance, while ensuring benefit for the ecosystems within these habitats.

HOW DO MAGELLANIC PENGUINS COPE WITH VARIABILITY IN THEIR ACCESS TO PREY

Movements of animals provisioning offspring by central place foraging extend from short, highly local trips where food is brought back essentially unchanged from its normal condition to extensive interseasonal movement where the offspring are nourished from body reserves built up during the adults absence from the breeding site. Here, appropriate strategies for maximizing lifetime reproductive success depend on the abundance and location of prey in relation to breeding sites and the energetics and speed of travel of the animal. Magellanic Penguins Spheniscus magellanicus undertake central place movements that are particularly variable during the incubation period; trips may last from a single day to over three weeks depending on colony locality. We reasoned that site-specific variability in prey distribution and abundance is responsible for this. Remote-sensing systems attached to 92 penguins from six different colonies over the species distributional range over the Patagonian Shelf were used to determine space use and foraging patterns in an attempt to understand the observed patterns. Birds in the north and south of the latitudinal range were essentially monophagic, feeding primarily on anchovies Engraulis anchoita and sprats Sprattus fuegensis, respectively, both species that are to be found relatively close to the colonies. Penguins in the center of the distributional range, where these pelagic school fish prey are essentially absent at that time of the year, traveled either north or south, to the same regions utilized by their conspecifics, presumably to exploit the same prey. A simple model is used to clarify patterns and can be used to predict which movement strategy is likely to be best according to colony location. During chick rearing, southerly movement of anchovies and northerly movement of sprats mean that Magellanic Penguins in the center of the distributional range may benefit, although the abundance of these fish is considered to be less than that closer to the Magellanic Penguin range limits. The extensive time involved in the foraging trips during incubation coupled with the postulated poorer prey conditions during the chick-rearing phase may help explain why Magellanic Penguin colony sizes in the center of the range are not elevated.

Fine‐scale foraging behaviour of a medium‐ranging marine predator

1. Movement patterns of predators should allow them to detect and respond to prey patches at different spatial scales, particularly through the adoption of area-restricted search (ARS) behaviour. Here we use fine-scale movement and activity data combined with first-passage time (FPT) analysis to examine the foraging strategy of northern gannets Morus bassanus in the western North Sea, and to test the following hypotheses: (i) birds adopt a hierarchical foraging strategy characterized by nested ARS behaviour; (ii) the locations and characteristics of ARS zones are strongly influenced by physical oceanography; (iii) the initiation of ARS behaviour is triggered by the detection and pursuit of prey; (iv) ARS behaviour is strongly linked to increased foraging effort, particularly within nested ARS areas. 2. Birds on 13 of 15 foraging trips adopted ARS behaviour at a scale of 9.1 +/- 1.9 km, and birds on 10 of these 13 trips adopted a second, nested ARS scale of 1.5 +/- 0.8 km, supporting hypothesis 1 above. ARS zones were located 117 +/- 55 km from the colony and over half were within 5 km of a tidal mixing front ~50 km offshore, supporting hypothesis 2 above. 3. The initiation of ARS behaviour was usually followed after only a short time interval (typically ~5 min) by the commencement of diving. Gannets do not dive until after they have located prey, and so this pattern strongly suggests that ARS behaviour was triggered by prey detection, supporting hypothesis 3 above. However, ~33% of dives in mixed coastal water and 16% of dives in stratified water were not associated with any detectable ARS behaviour. Hence, while ARS behaviour resulted from the detection and pursuit of prey, encounters with prey species did not inevitably induce ARS behaviour. 4. Following the initiation of ARS behaviour, dive rates were almost four times higher within ARS zones than elsewhere and almost three times higher in zones with nested ARS behaviour than in those without, supporting hypothesis 4 above and suggesting that the foraging success of birds was linked to their ability to match the hierarchical distribution of prey.

Using behavioural and state variables to identify proximate causes of population change in a seabird

Changes in animal population size are driven by the interactions between intrinsic processes and extrinsic forces, and identifying the proximate mechanisms behind population change remains a fundamental question in ecology. Here we report on how measuring behavioural and state proxies of food availability among populations experiencing different growth rates can be used to rapidly identify proximate drivers of population trends. In recent decades, the Cape gannet Morus capensis has shown a major distributional shift with historically large colonies in Namibia decreasing rapidly, whilst numbers at South African colonies have increased, suggesting contrasting environmental conditions in the two regions. We compared per capita growth rates of five of the six extant colonies with foraging range (using miniaturised Global Positioning System loggers), foraging work rate, food delivery rates and body condition of breeding adults. We found significant associations between the rate of population change, individual behaviour, energetic gain and body condition that indicate that recent population changes are associated with extrinsic effects. This study shows that behavioural and state data can be used to identify important drivers of population change, and their cost-effectiveness ensures that they are an appealing option for measuring the health of animal populations in numerous situations.

Shifts in phenotypic plasticity constrain the value of seabirds as ecological indicators of marine ecosystems

Marine ecosystems are critically challenged by human activities, urgently calling for better management practices. It has been proposed that conspicuous top predators such as seabirds may be used as ecological indicators. This approach requires intimate knowledge of relationships connecting seabird parameters to other ecosystem components (i.e., population plasticity, underlined by individual reaction norms), information which remains scarce. Furthermore, if seabirds are to be used as long-term indicators, the strength of the average plastic response in a studied population has to be sustained through time and space. This second aspect has so far been startlingly neglected, although previous studies underline shifts in the plasticity of seabird traits and detail the tools allowing an evolutionary and ecological study of plasticity in bird populations. Building upon these advances, we argue that gradual or sudden spatiotemporal changes in seabird phenotypic plasticity should not be neglected when designing monitoring schemes. We conclude that seabirds are best used as qualitative sentinels, rather than as quantitative indicators.

Adverse foraging conditions may impact body mass and survival of a high Arctic seabird

Tradeoffs between current reproduction and future survival are widely recognized, but may only occur when food is limited: when foraging conditions are favorable, parents may be able to reproduce without compromising their own survival. We investigated these tradeoffs in the little auk (Alle alle), a small seabird with a single-egg clutch. During 2005–2007, we examined the relationship between body mass and survival of birds breeding under contrasting foraging conditions at two Arctic colonies. We used corticosterone levels of breeding adults as a physiological indicator of the foraging conditions they encountered during each reproductive season. We found that when foraging conditions were relatively poor (as reflected in elevated levels of corticosterone), parents ended the reproductive season with low body mass and suffered increased post-breeding mortality. A positive relationship between body mass and post-breeding survival was found in one study year; light birds incurred higher survival costs than heavy birds. The results of this study suggest that reproducing under poor foraging conditions may affect the post-breeding survival of long-lived little auks. They also have important demographic implications because even a small change in adult survival may have a large effect on populations of long-lived species.

Cormorants dive through the Polar night

Most seabirds are visual hunters and are thus strongly affected by light levels. Dependence on vision should be problematic for species wintering at high latitudes, as they face very low light levels for extended periods during the Polar night. We examined the foraging rhythms of male great cormorants (Phalacrocorax carbo) wintering north of the Polar circle in West Greenland, conducting the first year-round recordings of the diving activity in a seabird wintering at high latitudes. Dive depth data revealed that birds dived every day during the Arctic winter and did not adjust their foraging rhythms to varying day length. Therefore, a significant proportion of the dive bouts were conducted in the dark (less than 1 lux) during the Polar night. Our study underlines the stunning adaptability of great cormorants and raises questions about the capacity of diving birds to use non-visual cues to target fish.

Black and white under the South African sun: are juvenile Cape gannets heat stressed?

(1) We measured dorsal surface temperature (Td), skin temperature (Ts) and body temperature (Tb) of adult (white plumage) and juvenile (black plumage) Cape gannets, Morus capensis, subject to high ambient temperatures. (2) Mean Td and Tb of juveniles were significantly elevated compared to those of adults. (3) Mean Ts of older juveniles, which had down beneath their body feathers, were significantly lower than Ts of younger juveniles, indicating that the down provided insulation. (4) Juveniles spent a significantly greater proportion of time thermoregulating via evaporative cooling than adults. (5) The high thermal load of the dark plumage is presumably associated with major costs for the thermal and water balance of juveniles.

Diving behaviour of African penguins: Do they differ from other Spheniscus penguins?

African penguins Spheniscus demersus closely resemble Magellanic S. magellanicus and Humboldt S. humboldti penguins and have similar breeding and feeding ecologies. Adults feed on pelagic schooling fish in continental shelf waters, but African penguins have been reported to have shallower dive angles and remain submerged longer for dives to a given depth than their congeners. The few data for African penguins were gathered using relatively large time-depth recorders. We measured diving behaviour of 36 African penguins provisioning small chicks at three colonies near Cape Town, South Africa. Maximum and mean dive depths were 69m and 14m respectively. Diving took place mainly during the day. Although dive depths differed between colonies, there were no significant differences in dive duration or maximum, median or mean depth. Total dive duration, descent time, bottom time, ascent time and dive angle all were strongly correlated with the maximum depth attained. The diving behaviour of African penguins is similar to that of its congeners. Diving performance probably was compromised by the data-logger used in the previous study. Comparative data from Humboldt penguins also indicate potential biases in an earlier study of this species. Care is needed when comparing the diving performance of penguins measured using different loggers.