Els M. van der Zee

Spatially extended habitat modification by intertidal reef-building bivalves has implications for consumer-resource interactions

Ecosystem engineers can strongly modify habitat structure and resource availability across space. In theory, this should alter the spatial distributions of trophically interacting species. In this article, we empirically investigated the importance of spatially extended habitat modification by reef-building bivalves in explaining the distribution of four avian predators and their benthic prey in the Wadden Sea—one of the world’s largest intertidal soft-sediment ecosystems. We applied Structural Equation Modeling to identify important direct and indirect interactions between the different components of the system. We found strong spatial gradients in sediment properties into the surrounding area of mixed blue mussel (Mytilus edulis) and Pacific oyster (Crassostrea gigas) reefs, indicating large-scale (100s of m) engineering effects. The benthic community was significantly affected by these gradients, with the abundance of several important invertebrate prey species increasing with sediment organic matter and decreasing with distance to the reefs. Distance from the reef, sediment properties, and benthic food abundance simultaneously explained significant parts of the distribution of oystercatchers (Haematopus ostralegus), Eurasian curlews (Numenius arquata), and bar-tailed godwits (Limosa lapponica). The distribution of black-headed gulls (Chroicocephalus ridibundus)—a versatile species with many diet options—appeared unaffected by the reefs. These results suggest that intertidal reef builders can affect consumer-resource dynamics far beyond their own boundaries, emphasizing their importance in intertidal soft-bottom ecosystems like the Wadden Sea.

Cross-habitat interactions among bivalve species control community structure on intertidal flats

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

Territory quality affects female preference in a Lake Victoria cichlid fish

Mate preferences on male colour have been implicated in generating and maintaining species diversity among haplochromine cichlid fish. Their lek-like mating system suggests that not only male colour but also territory quality is instrumental in mate choice. We assessed the relative importance of territory quality and male colour in mate choice by testing whether territory quality can override the female preference for males of her own colour in the Lake Victoria cichlid genus Pundamilia. First, we showed in experimental groups that the dominant male preferentially monopolised a large tube relative to a small tube. The situation mimics quality difference in rocky crevices that serves as a focal point for male courtship display. Second, in mate-choice tests, Pundamilia nyererei females were allowed to choose between closely related P. nyererei and P. pundamilia males; these species differ strikingly in male nuptial coloration, but little else. We gave either both males the same small tube or one of them a large tube. The preference of P. nyererei females for P. nyererei males in the control situation (where both males had a small tube) was significantly diminished in favour of P. pundamilia males when the latter had the large tube. The results provide experimental evidence that differences in territory quality can override the female preference for males of her own colour. This finding is critical for a recent hypothesis proposing that male competition for mating territories can facilitate the process of sympatric speciation by sexual selection.

Ecosystem engineering by seagrasses interacts with grazing to shape an intertidal landscape

Self-facilitation through ecosystem engineering (i.e., organism modification of the abiotic environment) and consumer-resource interactions are both major determinants of spatial patchiness in ecosystems. However, interactive effects of these two mechanisms on spatial complexity have not been extensively studied. We investigated the mechanisms underlying a spatial mosaic of low-tide exposed hummocks and waterlogged hollows on an intertidal mudflat in the Wadden Sea dominated by the seagrass Zostera noltii. A combination of field measurements, an experiment and a spatially explicit model indicated that the mosaic resulted from localized sediment accretion by seagrass followed by selective waterfowl grazing. Hollows were bare in winter, but were rapidly colonized by seagrass during the growth season. Colonized hollows were heavily grazed by brent geese and widgeon in autumn, converting these patches to a bare state again and disrupting sediment accretion by seagrass. In contrast, hummocks were covered by seagrass throughout the year and were rarely grazed, most likely because the waterfowl were not able to employ their preferred but water requiring feeding strategy (‘dabbling’) here. Our study exemplifies that interactions between ecosystem engineering by a foundation species (seagrass) and consumption (waterfowl grazing) can increase spatial complexity at the landscape level.

Habitat-Mediated Facilitation and Counteracting Ecosystem Engineering Interactively Influence Ecosystem Responses to Disturbance

Recovery of an ecosystem following disturbance can be severely hampered or even shift altogether when a point disturbance exceeds a certain spatial threshold. Such scale-dependent dynamics may be caused by preemptive competition, but may also result from diminished self-facilitation due to weakened ecosystem engineering. Moreover, disturbance can facilitate colonization by engineering species that alter abiotic conditions in ways that exacerbate stress on the original species. Consequently, establishment of such counteracting engineers might reduce the spatial threshold for the disturbance, by effectively slowing recovery and increasing the risk for ecosystem shifts to alternative states. We tested these predictions in an intertidal mudflat characterized by a two-state mosaic of hummocks (humps exposed during low tide) dominated by the sediment-stabilizing seagrass Zostera noltii) and hollows (low-tide waterlogged depressions dominated by the bioturbating lugworm Arenicola marina). In contrast to expectations, seagrass recolonized both natural and experimental clearings via lateral expansion and seemed unaffected by both clearing size and lugworm addition. Near the end of the growth season, however, an additional disturbance (most likely waterfowl grazing and/or strong hydrodynamics) selectively impacted recolonizing seagrass in the largest (1 m2) clearings (regardless of lugworm addition), and in those medium (0.25 m2) clearings where lugworms had been added nearly five months earlier. Further analyses showed that the risk for the disturbance increased with hollow size, with a threshold of 0.24 m2. Hollows of that size were caused by seagrass removal alone in the largest clearings, and by a weaker seagrass removal effect exacerbated by lugworm bioturbation in the medium clearings. Consequently, a sufficiently large disturbance increased the vulnerability of recolonizing seagrass to additional disturbance by weakening seagrass engineering effects (sediment stabilization). Meanwhile, the counteracting ecosystem engineering (lugworm bioturbation) reduced that threshold size. Therefore, scale-dependent interactions between habitat-mediated facilitation, competition and disturbance seem to maintain the spatial two-state mosaic in this ecosystem.

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.

Copper treatment during storage reduces Phytophthora and Halophytophthora infection of Zostera marina seeds used for restoration

Restoration is increasingly considered an essential tool to halt and reverse the rapid decline of vital coastal ecosystems dominated by habitat-forming foundation species such as seagrasses. However, two recently discovered pathogens of marine plants, Phytophthora gemini and Halophytophthora sp. Zostera, can seriously hamper restoration efforts by dramatically reducing seed germination. Here, we report on a novel method that strongly reduces Phytophthora and Halophytophthora infection of eelgrass (Zostera marina) seeds. Seeds were stored in seawater with three different copper sulphate concentrations (0.0, 0.2, 2.0 ppm) crossed with three salinities (0.5, 10.0, 25.0 ppt). Next to reducing seed germination, infection significantly affected cotyledon colour: 90% of the germinated infected seeds displayed a brown cotyledon upon germination that did not continue development into the seedling stage, in contrast to only 13% of the germinated non-infected seeds. Copper successfully reduced infection up to 86% and the 0.2 ppm copper sulphate treatment was just as successful as the 2.0 ppm treatment. Infection was completely eliminated at low salinities, but green seed germination was also dramatically lowered by 10 times. We conclude that copper sulphate treatment is a suitable treatment for disinfecting Phytophthora or Halophytophthora infected eelgrass seeds, thereby potentially enhancing seed-based restoration success.

Banc d'Arguin - Food webs with trophic effects by seagrass removed

This file contains matrices from which we we removed seagrass (Zostra noltii) and any species exclusively feeding on it.