An evolutionary perspective on caching by corvids

A principal finding in the food-caching literature is that species differences in hoarding propensity are positively correlated with species differences in degree of adaptations to caching behaviour, such as performance on spatial memory tasks and hippocampal volume. However, there are examples that do not fit this pattern. We argue that these examples can be better understood by considering the phylogenetic relatedness between species. We reconstruct the ancestral state for caching behaviour in corvids and assess when transitions in caching behaviour occurred within the corvid phylogeny. Our analysis shows that the common ancestor of all corvids was a moderate cacher. This result suggests that corvids followed a bi-directional evolutionary trajectory in which caching was secondarily lost twice and there were at least two independent transitions from moderate to specialized caching. The independent evolution of specialized cachers in the two groups must, therefore, be a case of convergent evolution. This is exemplified by the fact that specialized cachers show structurally different adaptations serving the same function to intense caching, such as different pouches to transport food. Finally, we argue that convergent evolution may have led to adaptations in memory and hippocampus that serve the same function but differ in design, and that these different adaptations may explain the examples that do not fit the pattern predicted by the adaptive specialization hypothesis

Estimating the contribution of carnivorous waterbirds to nutrient loading in freshwater habitats

1. We estimated nitrogen (N) and phosphorus (P) loading into wetlands by carnivorous waterbirds with alternative physiological models using a food-intake and an excreta-production approach. The models were applied for non-breeding and breeding Dutch inland carnivorous waterbird populations to quantify their contribution to nutrient loading on a landscape scale. 2. Model predictions based on food intake exceeded those based on excretion by 59–62% for N and by 2–36% for P, depending on dietary assumptions. Uncertainty analysis indicated that the intake model was most affected by errors in energy requirement, while the excretion model was dependent on faecal nutrient composition. 3. Per capita loading rate of non-breeders increased with body mass from 0.3–0.8 g N day1 and 0.15 g P day1 in little gulls Larus minutus to 4.5–11.5 g N day1 and 2.1–3.2 g P day1 in great cormorants Phalacrocorax carbo. For breeding birds, the estimated nutrient loading by a family unit over the entire breeding period ranged between 17.6–443.0 g N and 8.6 g P for little tern Sterna albifrons to 619.6–1755.6 g N and 316.2–498.1 g P for great cormorants. 4. We distinguished between external (i.e. importing) and internal (i.e. recycling) nutrient loading by carnivorous waterbirds. For the Netherlands, average external-loading estimates ranged between 38.1–91.5 tonnes N and 16.7–18.2 tonnes P per year, whilst internal-loading estimates ranged between 53.1–140.5 tonnes N and 25.2–39.2 tonnes P and per year. The average contribution of breeding birds was estimated to be 17% and 32% for external and internal loading respectively. Most important speci 5. On a landscape scale, loading by carnivorous waterbirds was of minor importance for freshwater habitats in the Netherlands with 0.26–0.65 kg N ha1 a1 and 0.12–0.16 kg P ha1 a1. However, on a local scale, breeding colonies may be responsible for significant P loading.