Does logging and forest conversion to oil palm agriculture alter functional diversity in a biodiversity hotspot?

Forests in Southeast Asia are rapidly being logged and converted to oil palm. These changes in land-use are known to affect species diversity but consequences for the functional diversity of species assemblages are poorly understood. Environmental filtering of species with similar traits could lead to disproportionate reductions in trait diversity in degraded habitats. Here, we focus on dung beetles, which play a key role in ecosystem processes such as nutrient recycling and seed dispersal. We use morphological and behavioural traits to calculate a variety of functional diversity measures across a gradient of disturbance from primary forest through intensively logged forest to oil palm. Logging caused significant shifts in community composition but had very little effect on functional diversity, even after a repeated timber harvest. These data provide evidence for functional redundancy of dung beetles within primary forest and emphasize the high value of logged forests as refugia for biodiversity. In contrast, conversion of forest to oil palm greatly reduced taxonomic and functional diversity, with a marked decrease in the abundance of nocturnal foragers, a higher proportion of species with small body sizes and the complete loss of telecoprid species (dung-rollers), all indicating a decrease in the functional capacity of dung beetles within plantations. These changes also highlight the vulnerability of community functioning within logged forests in the event of further environmental degradation.

Influence of wing loading on the trade-off between pursuit-diving and flight in common guillemots and razorbills.

Species of bird that use their wings for underwater propulsion are thought to face evolutionary trade-offs between flight and diving, leading to the prediction that species with different wing areas relative to body mass (i.e. different wing loadings) also differ in the relative importance of flight and diving activity during foraging trips. We tested this hypothesis for two similarly sized species of Alcidae (common guillemots and razorbills) by using bird-borne devices to examine three-dimensional foraging behaviour at a single colony. Guillemots have 30% higher wing loading than razorbills and, in keeping with this difference, razorbills spent twice as long in flight as a proportion of trip duration whereas guillemots spent twice as long in diving activity. Razorbills made a large number of short, relatively shallow dives and spent little time in the bottom phase of the dive whereas guillemots made fewer dives but frequently attained depths suggesting that they were near the seabed (ca. 35-70 m). The bottom phase of dives by guillemots was relatively long, indicating that they spent considerable time searching for and pursuing prey. Guillemots also spent a greater proportion of each dive bout underwater and had faster rates of descent, indicating that they were more adept at maximising time for pursuit and capture of prey. These differences in foraging behaviour may partly reflect guillemots feeding their chicks single large prey obtained near the bottom and razorbills feeding their chicks multiple prey from the water column. Nonetheless, our data support the notion that interspecific differences in wing loadings of auks reflect an evolutionary trade-off between aerial and underwater locomotion.

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.

Modelling the spatial distribution of ammonia emissions from seabirds in the UK.

Knowledge of the sources and distribution of ammonia (NH3) emissions underpins our understanding of the nitrogen budget. Research has focused on quantifying NH3 emissions from anthropogenic sources, whilst those from natural sources have received little attention internationally. Seabirds excrete large quantities of nitrogen, making seabird colonies a major natural source of NH3. Ammonia emissions from each UK seabird species were estimated and combined with population distribution data to model their spatial distribution. Total NH3 emissions from UK seabirds were estimated at 2.7 kt per year. Seabird emissions are concentrated in remote parts of the UK where anthropogenic emissions are small, so that seabirds often represent the main source of NH3 emissions in these areas. Seabird NH3 emissions were found to have increased by 34% since the 1970s. This corresponds to population changes which may be influenced by human activities, showing that even this natural source can be anthropogenically modified.

Sex-specific foraging behaviour in a monomorphic seabird.

Sexual differences in the foraging behaviour of parents have been observed in a number of sexually sizedimorphic birds, particularly seabirds, and the usual inference has been that these sex-specific differences are mediated primarily by differences in body size. To test this explanation, we compared the foraging behaviour of parents in a monomorphic seabird species, the northern gannet Morus bassanus. Using specially designed instruments and radio telemetry we found that individuals of both sexes were consistent in the directions and durations of their foraging trips. However, there were significant differences in the foraging behaviour of males and females. Female gannets were not only more selective than males in the areas where they foraged, but they also made longer, deeper dives and spent more time on the sea surface than males. As the sexes are morphologically similar in this species, then these differences are unlikely to have been mediated by body size. Our work highlights the need to investigate sexual differences in the foraging behaviour of seabirds and other species more closely, in order to test alternative theories that do not rely on differences in body size.

Evidence of intra-specific competition for food in a pelagic seabird.

The factors affecting the population dynamics of seabirds have long intrigued biologists. Current data suggest that density-dependent depletion of prey during the breeding season may regulate population size. However, much of the evidence for this has been circumstantial, and the underlying mechanisms are unclear. Here, we show that the per capita population growth rates of northern gannet Morus bassanus at colonies in Britain and Ireland have declined with increasing population size. Furthermore, direct observations reveal that the mean foraging trip duration of breeding gannets is positively correlated with colony size, both among colonies of different sizes in the same year, and within colonies as they change in size. To understand this phenomenon, we have developed a model which demonstrates that disturbance of fish alone can readily generate conditions under which gannets at larger colonies have to travel further to obtain food.

Postnatal development of northern fulmar chicks, Fulmarus glacialis.

The slow growth and large fat stores characteristic of many pelagic seabird chicks were generally assumed to reflect infrequent and unpredictable food provisioning by parents. Much less attention has been focused on the importance of intrinsic physiological processes in shaping patterns of development. In this study, we examined postnatal growth and changes in water content of different organs in fulmar chicks, Fulmarus glacialis, from Fair Isle, United Kingdom. After correcting for body size, mass growth rate was as high as in inshore-feeding species, which did not support the notion of an external constraint on growth imposed by the unpredictability of pelagic prey. Pectoral muscles and plumage grew more rapidly than other tissues. Pectorals also had a high water index, probably indicating slower maturation compared with leg muscles, which need to generate heat earlier on to free adults from brooding requirements. Lean dry mass of liver, kidney, and gut decreased markedly toward fledging, presumably because of high energetic costs of maintaining large metabolic machinery in older chicks and analogous to the situation in adult waders before migration. These results suggest that the general pattern of development of fulmars may be linked to changes in resource allocation as chicks grow and possibly a compromise at the tissue level between cell division and the attainment of mature function.

The interactive effects of age and food supply on the breeding ecology of great skuas.

Age-specific variation in the reproductive performance of great skuas in Shetland, U.K., was investigated during a period of fluctuating sandeel availability between 1988 and 1993. Increased sandeel abundance was associated with earlier laying dates, increased clutch volumes and improved fledging success. Sandeel abundance had no effects on clutch size and hatching success. Parental age improved haying date and clutch size in a nonlinear manner. Laying date became earlier in the youngest age classes, with the effect being negligible in older birds. Clutch size increased with age up to 18 and then declined in older birds. Clutch volume increased with age and the probability of nest predation declined with age. There were no significant interactive effects of age and year on laying date nor clutch volume, indicating that birds of all ages benefited equally from increases in sandeel abundance. This suggests that lack of breeding experience rather than deficient foraging skills are responsible for young birds laying later. There was a significant interactive effect of year and age on fledging success that was associated with changes in sandeel availability. Success was uniformly low for birds of all ages in 1988-90 when sandeel abundance was low. Success increased with age in 1991 and 1992 during which time food supply improved. During 1993 food supply was abundant and fledging success was high in all age classes. Age-specific improvements in laying date, clutch size and nest predation were best explained by learning of skills associated with experience of breeding. Improvements in fledging success with age were probably best explained by the learning of skills associated with foraging.

Nestling obesity in procellariiform seabirds: temporal and stochastic variation in provisioning and growth of short-tailed shearwaters Puffinus tenuirostris.

Procellariiform seabirds such as short-tailed shearwaters Puffinus tenuirostris accumulate large quantities of lipid during the nestling period. The functional significance of this pattern of development remains unclear, but has been related both to temporal variation in feeding conditions around the colony and to stochastic variation in the foraging success of individual parents. This paper examines temporal and age-specific variation in the pattern of food delivery to nestling short-tailed shearwaters, which have one of the lowest provisioning rates of any procellariiforms and are known to experience occasional long intervals between feeds. We assess whether variation in the provisioning rates of chicks was associated primarily with temporal variation in food delivery at the level of the colony or with stochastic variation in food delivery at the level of the individual. We then discuss this variability in the context of nestling obesity. For all but the youngest chick age-classes, individual meals delivered by adults averaged 141 g, which was 25% of adult body mass. The proportion of chicks fed each night was low (49%) and highly variable (coefficient of variation = 82%), which means that occasional long intervals between feeds would be expected to arise simply by chance. In keeping with this, intervals between feeding events for individual chicks followed a negative exponential distribution with a mean of 2 nights and a maximum interval of 13 nights. There was significant temporal variation in food delivery, but deviations from expected values for both feeding frequency and meal size were restricted to a small number of nights, included values both higher and lower than expected and did not persist for more than 2 nights in succession. These data suggest that even among those species with very low feeding frequencies and occasional long intervals between feeds, nestling obesity in Procellariiformes should be regarded as a response to chronic stochastic variability in food delivery at the level of the individual chick rather than as insurance against sporadic temporal variation at the level of the colony.