Sander Holthuijsen

Burrowing behavior of a deposit feeding bivalve predicts change in intertidal ecosystem state

Behavior has a predictive power that is often underutilized as a tool for signaling ecological change. The burrowing behavior of the deposit feeding bivalve Macoma balthica reflects a typical food-safety trade-off. The choice to live close to the sediment surface comes at a risk of predation and is a decision made when predation danger, food intake rates or future fitness prospects are low. In parts of the Dutch Wadden Sea, Macoma populations declined by 90% in the late 1990s, in parallel with large-scale mechanical cockle-dredging activities. During this decline, the burrowing depth of Macoma became shallow and was correlated with the population decline in the following year, indicating that it forecasted population change. Recently, there has been a series of large recruitment events in Macoma. According to the food-safety trade-off, we expected that Macoma should now live deeper, and have a higher body condition in association with this change in depth of living. Indeed, we observed that Macoma now lives deeper and that living depth in a given year forecasted population growth to the next year, especially in individuals larger than 14 mm. As living depth and body condition were strongly correlated in individuals larger than 14 mm, larger Macoma could be living deeper to protect their reproductive assets. Our results confirmed that burrowing depth signals impending population change and, together with body condition, can provide an early warning signal of ecological change. We suggest that population recovery is being driven by improved intertidal habitat quality in the Dutch Wadden Sea, rather than by the proposed climate-change related effects. This shift in ecosystem state is suggested to include the recovery of diatom habitat in the top layer of the sediment after cockle-dredging ended.

Understanding spatial distributions: negative density-dependence in prey causes predators to trade-off prey quantity with quality

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The ‘functional response’ couples a predators intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.

Presence-absence of marine macrozoobenthos does not generally predict abundance and biomass

Many monitoring programmes of species abundance and biomass increasingly face financial pressures. Occupancy is often easier and cheaper to measure than abundance or biomass. We, therefore, explored whether measuring occupancy is a viable alternative to measuring abundance and biomass. Abundance- or biomass-occupancy relationships were studied for sixteen macrozoobenthos species collected across the entire Dutch Wadden Sea in eight consecutive summers. Because the form and strength of these relationships are scale-dependent, the analysis was completed at different spatiotemporal scales. Large differences in intercept and slope of abundance- or biomass-occupancy relationships were found. Abundance, not biomass, was generally positively correlated with occupancy. Only at the largest scale, seven species showed reasonably strong abundance-occupancy relationships with large coefficients of determination and small differences in observed and predicted values (RMSE). Otherwise, and at all the other scales, intraspecific abundance and biomass relationships were poor. Our results showed that there is no generic relationship between a species’ abundance or biomass and its occupancy. We discuss how ecological differences between species could cause such large variation in these relationships. Future technologies might allow estimating a species’ abundance or biomass directly from eDNA sampling data, but for now, we need to rely on traditional sampling technology.