Why do Gull-billed Terns Gelochelidon nilotica feed on fiddler crabs Uca tangeri in Guinea-Bissau?

Gull-billed Terns Gelochelidon nilotica wintering in Guinea Bissau mainly fed on fiddler crabs Uca tangeri and were occasionally seen feeding on fish and locusts. As fiddler crabs have a low energy content, terns need a large gross intake to meet daily energy demands. Fiddler crabs also have a low ratio of digestible flesh to exoskeleton, and therefore tern food intake may be limited by gut capacity. Activity budgets of Gullbilled Terns feeding on fiddler crabs showed that a considerable part of the time was spent resting. The duration of resting intervals increased with energy intake and was positively correlated with the metabolisable energy content of the crab eaten, suggesting that resting periods were required for a proper digestion. The poor quality of fiddler crabs was offset by high capture rates. So daily energy expenditure of the terns could easily be met by feeding on fiddler crabs. Even when resting pauses were included in foraging time, foraging for only 1.5 hours on fiddler crabs satisfied the terns’ daily energy demands. Instead, feeding on energy-rich fish would require about 2.5 hours to satisfy daily energy demands. Compared to the more specialised piscivorous Little Tern Sternula albifrons and Sandwich Tern Sterna sandvicensis, capture rate of fish was poor in Gull-billed Terns. From an energetic point of view, wintering Gull-billed Terns feeding on fiddler crabs seem to have an easy living in Guinea Bissau.

Flocking and feeding in the fiddler crab (UCA tangeri): prey availability as risk-taking behaviour

For a full understanding of prey availability, it is necessary to study risk-taking behaviour of the prey. Fiddler crabs are ideally suited for such a study, as they have to leave their safe burrow to feed on the surface of the intertidal flats during low tide, thereby exposing themselves to avian predators. A study in an intertidal area along the coast of Mauritania showed that small crabs always stayed in the vicinity of their burrow, but large crabs wandered in large flocks (also referred to as droves) to feed on sea-grass beds downshore. Transplanting downshore feeding substrate to the burrowing zone of the small crabs proved that they too preferred to feed on it. Since small crabs can be preyed upon by more species of birds, this suggests that the decision not to leave the burrowing zone might be related to the risk of being fed upon by birds. We calculated predation risk from measurements on the density and feeding activity of the crabs, as well as the feeding density, the intake rate and the size selection of the avian predators. Per hour on the surface, crabs in a flock were more at risk than crabs feeding near their burrow. Thus, though flocking crabs may have benefited from ‘swamping the predator’ by emerging in maximum numbers during some tides only, this did not reduce their risk of predation below that of non-flocking crabs. Furthermore we found that irrespective of activity, large crabs suffered a higher mortality per tide from avian predators than small crabs. This suggests that large crabs could not sufficiently reduce their foraging time to compensate for the increased risk while foraging in a flock, even though they probably experienced better feeding conditions than small crabs staying near their burrow. The greater energy demands of large crabs were reflected in a greater surface area grazed. Thus, with increasing size a fiddler crab has to feed further away from its burrow and so may derive less protection from staying near to it. It seems that growing big does not reduce the risk of predation for fiddler crabs, as it does in many other species with indeterminate growth. As in such species, the most probable advantage of growing big is increased mating success. Ultimately, therefore, prey availability must be understood from the life-history decisions of the prey species.

Habitat selection and energetics of the fiddler crab (Uca tangeri)

We tried to unravel the possible links between the skewed predation risk in Uca tangeri (where large individuals are more at risk from avian predators) and size-dependent changes in the physiology and habitat choice of this fiddler crab species. Over a transect running from low to high in the tidal zone of a beach in Mauritania, the temperature profile at various depths in the substrate, the water-table level of seep water, salt concentration of seep water, depth of the aerobic level, operative temperatures on the surface, and size distribution of crabs were assessed. In addition, resting metabolic rates, Q10 and thermal and starvation tolerances were estimated. Going from low to high in the tidal zone, crab size and burrow depth increased. At the preferred burrowing depth, microclimatological conditions appeared to be equally favourable at all sites. At the surface, conditions were more favourable low in the tidal zone, where also food availability is sufficient to enable small crabs to forage in the vicinity of their burrows. Large crabs have higher energy requirements and are thereby forced to forage in flocks low in the tidal zone where food is probably more abundant. Low in the tidal zone, digging deeply is impossible as the aerobic layer is rather thin. Large crabs prefer living high in the tidal zone as (1) deep burrows ensure better protection against predators, (2) more time is available for digging holes and (3) the substrate is better suited for reproduction. Energy reserves in late summer ensured an average of 34 days of survival. It is argued that the allotment of energy to growth must be considerable even in reproducing animals; the rewards of growth being the disproportional increase in reproductive output with size.