Bonamia exitiosa transmission among, and incidence in, Asian oyster Crassostrea ariakensis under warm euhaline conditions.

Previously reported in Australia, New Zealand, and more recently in Europe, the protistan parasite Bonamia exitiosa was also reported in the mid-Atlantic region of the USA after causing serious mortalities there in the Asian oyster Crassostrea ariakensis. At the time, this oyster was being considered for introduction, and the potential consequences of introducing this species were being assessed using field and laboratory studies. B. exitiosa emerged as the most serious disease threat for this oyster species, especially under warm euhaline conditions and for oysters <50 mm in size. To better evaluate how quickly this parasite may be able to spread among C. ariakensis, we investigated B. exitiosa transmission and incidence in C. ariakensis. During a first trial, potential direct transmission of B. exitiosa was evaluated by cohabitating infected C. ariakensis with uninfected C. ariakensis under in vivo quarantine conditions. In a second experiment, B. exitiosa incidence was estimated in situ by determining its prevalence in C. ariakensis deployed in an enzootic area after 4, 7, 14, 21 and 28 d of exposure. Results suggest that under warm euhaline conditions B. exitiosa can be transmitted among C. ariakensis without requiring any other parasite source and that parasite incidence may be at least as high as 40% after only 4 d exposure to an enzootic area. These results underscored the severity of the bonamiasis disease threat to C. ariakensis and provided further evidence that efforts to build an aquaculture industry based on C. ariakensis in the eastern USA might have been thwarted by parasitic disease.

Historical overfishing and the recent collapse of coastal ecosystems.

Ecological extinction caused by overfishing precedes all other pervasive human disturbance to coastal ecosystems, including pollution, degradation of water quality, and anthropogenic climate change. Historical abundances of large consumer species were fantastically large in comparison with recent observations. Paleoecological, archaeological, and historical data show that time lags of decades to centuries occurred between the onset of overfishing and consequent changes in ecological communities, because unfished species of similar trophic level assumed the ecological roles of overfished species until they too were overfished or died of epidemic diseases related to overcrowding. Retrospective data not only help to clarify underlying causes and rates of ecological change, but they also demonstrate achievable goals for restoration and management of coastal ecosystems that could not even be contemplated based on the limited perspective of recent observations alone.

Tubeworm succession at hydrothermal vents: use of biogenic cues to reduce habitat selection error?

Species colonizing new deep-sea hydrothermal vents along the East Pacific Rise show a distinct successional sequence: pioneer assemblages dominated by the vestimentiferan tubeworm Tevnia jerichonana being subsequently invaded by another vestimentiferan Riftia pachyptila, and eventually the mussel Bathymodiolus thermophilus. Using a manipulative approach modified from shallow-water ecological studies, we test three alternative hypotheses to explain the initial colonization by T. jerichonana and its subsequent replacement by R. pachyptila. We show that R. pachyptila and another vestimentiferan, Oasisia alvinae, colonized new surfaces only if the surfaces also were colonized by T. jerichonana. This pattern does not appear to be due to restricted habitat tolerances or inferior dispersal capabilities of R. pachyptila and O. alvinae, and we argue the alternative explanation that T. jerichonana facilitates the settlement of the other two species and is eventually outcompeted by R. pachyptila. Unlike the classic model of community succession, in which facilitating species promote their own demise by modifying the environment to make it more hospitable for competitors, we suggest that T. jerichonana may produce a chemical substance that induces settlement of these competitors. This process of selecting habitat based on biogenic cues may be especially adaptive and widespread among later-successional species that occupy a physically variable and unpredictable environment. In these cases, the presence of weedy species implies some integrated period of environmental suitability, whereas an instantaneous assessment of physical habitat conditions, such as water temperature for vent tubeworms, provides a poorer predictor of long-term habitat suitability.

Modification of animal habitat by large plants: mechanisms by which seagrasses influence clam growth.

Field experiments withMercenaria mercenaria in a relatively high-energy environment demonstrated that clams on unvegetated sand flats failed to grow during autumn while those within seagrass beds grew substantially. Clam growth rates at the seagrass margin that first receives the faster-flowing, flood-tidal currents were about 25% less than at the opposite edge. In a second experiment, pruning, which reduced average blade length by 50-75%, was shown to enhance near-bottom current velocities and to reduce shell growth ofMercenaria during summer by about 50%. As in the first experiment, clams in the unvegetated sand flats exhibited no net growth. Clam mortality, caused mostly by predatory crabs and whelks, was much higher on sand flats than in seagrass beds and intermediate in clipped seagrass. Although consistent with some previous reports, these growth results are still surprising given that they contradict the generalization that suspension feeders grow faster under more rapid current regimes.Three types of indirect interactions might explain the observed effect of seagrass on growth of buried clams: (1) altering food supply; (2) changing the intensity of biological disturbance on feeding clams; and/or (3) affecting the physical stability of the sediments. Previous research on this question has focused almost exclusively on processes that alter food supply rates. In this study, food concentrations, as indicated by suspended chla, were 30% higher inside than outside one seagrass bed, whereas chla concentrations in two other beds were not different from those on adjacent sand flats. This result is sufficient to show that more intense food depletion was not induced by the reduction in flow velocities under the seagrass canopy. Nevertheless, the possible small difference in food concentrations between vegetated and unvegetated bottom seems insufficient to explain the absence of growth of sand-flat clams, especially given the virtual lack of food limitation among suspension feeders in this system. Two data sets demonstrated that the effects of biological disturbance agents cannot be ignored. An outdoor laboratory experiment showed that even in the absence of physical contact between predator and prey the presence of a whelk reduces the amount of time spent feeding byMercenaria. This result suggests that sand flats, where predation rates are higher, may be sites of lower clam growth than seagrass beds because of greater consumer interference with clam feeding. Furthermore, clam siphons are proportionately larger inside seagrass than on sand flats, implying that siphon nipping may not be as intense inside seagrass. This process, too, would reduce net growth of sand-flat clams. Finally, no explicit test was conducted of the hypothesis that enhanced sediment transport in the absence of flow baffling and root binding by seagrass inhibits net growth of clams on high-energy sand flats. Nevertheless, this is a reasonable explanation for the pattern of enhanced growth of seagrass clams, and could serve to explain the otherwise unexplained pattern of lower clam growth at the edge of the seagrass bed that experiences the faster flood-tidal current velocities. Each broad process, changing fluid dynamics, altering consumer access, and varying sediment stability, represents a mechanism whereby habitat structure, provided by the dominant plant, has an important indirect influence on the functional value of the habitat for resident animals.

Analysis of feeding preference experiments.

Published studies of consumer feeding preferences using foods that experience autogenic change in mass, numbers, area, etc., on the time scale of a feeding trial fail to employ appropriate statistical analyses to incorporate controls for those food changes occurring in the absence of the consumer. The studies that run controls typically use them to calculate a constant "correction factor", which is subtracted prior to formal data analysis. This procedure constitutes a non-rigorous suppression of variance that overstates the statistical significance of observed differences. The appropriate statistical analysis for preference tests with two foods is usually a simple t-test performed on the between-food differences in loss of mass (or numbers, area, etc.) comparing the results of experimentals with consumers to controls without consumers. Application of this recommended test procedure to an actual data set illustrates how low replication in controls, which is typical of most studies of feeding preference, inhibits detection of an apparently large influence of previous mechanical damage (simulated grazing) in reducing the attractiveness of a brown alga to a sea urchin.

Responses of growth to elevation fail to explain vertical zonation of suspension-feeding bivalves on a tidal flat.

Five species of suspension-feeding bivalves were transplanted to each of two elevations on a tidal flat at Shark Bay, Western Australia, at six replicate locations spaced at 1-km intervals along the shore. Four species exhibited greatly reduced growth at the higher elevation, while the fifth species did not respond to elevation. The magnitude of the % reductions in growth with increased elevation was 2-3 times the % reduction in average daily submergence, confirming a previous suggestion that differences in feeding time alone are insufficient to explain completely the reduced growth of suspension-feeding bivalves at higher tidal elevatios. All four species that responded showed the same pattern of higher growth lower on the shore, even though transect sampling showed that two were normally abundant only high on the shore while the other tow were naturally restricted to elevations low on the shore. Consequently, knowledge of how individual growth within species varies with tidal elevation fails to explain observed zonation patterns with elevation in this guild of suspension-feeding bivalves. The paradoxical distribution pattern of those two species that were rare at the lower tidal elevations, where they actually grew more rapidly, implies that some biological agent(s) of mortality not physiological stress set(s) their lower distributional limit on the shore. Biological rather than physical factors commonly, although not universally, set lower distributional limits of invertebrates in rocky intertidal zones, but this study provides the first experimental data to explore this concept in marine soft sediments.

Biological vs. physical explanations for the non-random pattern of host occupation by a macroalga attaching to infaunal bivalve molluscs.

On a shallow sand flat at Princess Royal Harbour near Albany, Western Australia, the brown macrophyte Hormosira banksii attaches to shells of infaunal bivalves. Hormosira occupies shells of Katelysia rhytiphora in preference to K. scalarina. We proposed and tested four hypotheses to explain this host occupation pattern. First, by following the fate of nearly one thousand marked clams of each species, we rejected the hypothesis that K. rhytiphora exhibits greater longevity and simply possesses more frequent Hormosira because of a longer temporal integration of settlement events. Second, we rejected the hypothesis that K. rhytiphora exhibits higher densities in the top 5 mm of sediment and accumulates more Hormosira because its shell is more abundant in the depth range occupied by the attaching base of Hormosira. Third, we showed that K. rhytiphora because of its larger size is more difficult to dislodge from the sediments than K. scalarina, supporting the hypothesis that Hormosira is rare on K. scalarina because storm waves selectively dislodge and carry to the beach Hormosira attached to K. scalarina. This physical explanation for the Hormosira occupation pattern gets further support from the observation that a third infaunal bivalve, the mussel Brachidontes erosus, has a far higher frequency of Hormosira occupation than either Katelysia species, while providing a much more robust anchor because of its extensive byssal attachments to neighboring mussels. The sizes of Hormosira plants also vary consistently with this physical transport hypothesis: Hormosira is smallest on K. scalarina and largest on B. erosus. Successful colonization of initially unoccupied Katelysia during five 3-9 month periods was also more frequent on K. rhytiphora than on K. scalarina. This suggests a fourth explanation for the Hormosira distribution pattern: that spore settlement is selective for K. rhytiphora in preference to K. scalarina. Although this hypothesis requires further testing, evolution of selective spore settlement would be reasonable given the different likelihoods of subsequent host dislodgement during storm waves.