Judi E. Hewitt

DISTURBANCE OF THE MARINE BENTHIC HABITAT BY COMMERCIAL FISHING: IMPACTS AT THE SCALE OF THE FISHERY

Commercial fishing is one of the most important human impacts on the marine benthic environment. One such impact is through disturbance to benthic habitats as fishing gear (trawls and dredges) are dragged across the seafloor. While the direct effects of such an impact on benthic communities appear obvious, the magnitude of the effects has been very difficult to evaluate. Experimental fishing-disturbance studies have dem- onstrated changes in small areas; however, the broader scale implications attributing these changes to fishing impacts are based on long-term data and have been considered equivocal. By testing a series of a priori predictions derived from the literature (mainly results of small-scale experiments), we attempted to identify changes in benthic communities at the regional scale that could be attributed to commercial fishing. Samples along a putative gradient of fishing pressure were collected from 18 sites in the Hauraki Gulf, New Zealand. These sites varied in water depth from -17 to 35 m and in sediment characteristics from -1 to 48% mud and from 3 to 8.5 (Lg chlorophyll a/cm3. Video transects were used for counting large epifauna and grab/suction dredge and core sampling were used for collecting macrofauna. After accounting for the effects of location and sediment characteristics, 15-20% of the variability in the macrofauna community com- position sampled in the cores and grab/suction dredge samples was attributed to fishing. With decreasing fishing pressure we observed increases in the density of echinoderms, long- lived surface dwellers, total number of species and individuals, and the Shannon-Weiner diversity index. In addition, there were decreases in the density of deposit feeders, small opportunists, and the ratio of small to large individuals of the infaunal heart urchin, Echino- cardium australe. The effects of fishing on the larger macrofauna collected from the grab/ suction dredge samples were not as clear. However, changes in the predicted direction in epifaunal density and the total number of individuals were demonstrated. As predicted, decreased fishing pressure significantly increased the density of large epifauna observed in video transects. Our data provide evidence of broad-scale changes in benthic communities that can be directly related to fishing. As these changes were identifiable over broad spatial scales they are likely to have important ramifications for ecosystem management and the development of sustainable fisheries.

Scale‐Dependent Recolonization: The Role of Sediment Stability in a Dynamic Sandflat Habitat

An important ecological issue is developing an understanding of how patterns and processes vary with scale. We designed a field experiment to test how differences in the aerial extent of disturbance affected macrofaunal recolonization on a sandflat. Three different plot sizes (0.203 m 2 , 0.81 m 2 , and 3.24 m 2 ) were defaunated, and samples were collected to assess recovery over a 9-mo period. As the sandflat used for the experiment was prone to disturbance by wind-driven waves, we also measured changes in sediment bed height (an indicator of sediment stability) over the course of the experiment. Most common species revealed significant relationships between density and disturbance plot size. Scale-dependent recovery was also demonstrated by differences in species assemblage structure over the course of the experiment. Relative rates of colonization varied by 50% between large and small experimental plots. However, these differences were not related to specific species, particular functional groups, or potential modes of colonization. The results revealed an unusually slow rate of faunal recovery following defaunation. Both increasing numbers of colonists and density changes in ambient sediments made an important contribution to recovery. The relationship found between changes in sediment bed height and wind velocity indicated that wind-driven wave disturbance was an important factor influencing sediment instability. Sediment instability was higher in all experimental plots than in the ambient sediments, due to the initial removal of a dense spionid polychaete tube mat characteristically found at the study site. Sediment instability also increased with increasing plot size. Thus in this dynamic sandflat habitat, faunal emigration from recovering disturbed patches of sediment may significantly slow rates of recolonization. These results demonstrate that incorporating patch size, emigration, recovery time, and interactions between hydrodynamic conditions and habitat stability (particularly where colonists influence sediment stability) are crucial to generating a general understanding of recovery processes in soft-sediment habitats. While our results demonstrate the need for caution in scaling-up from small-scale studies, they do indicate that larger scale disturbances that destroy organisms with a role in maintaining habitat stability are likely to result in very slow recovery dynamics, particularly in wave-disturbed soft-sediment habitats.

Muddy waters: elevating sediment input to coastal and estuarine habitats

Changes in land use and the development of coastal regions around the world have markedly increased rates of sediment input into estuarine and coastal habitats. Field studies looking at the consequences of terrestrial sediment deposition, water-borne sediment, and long-term changes in habitats indicate that increasing rates of sediment loading adversely affect the biodiversity and ecological value of estuarine and coastal ecosystems. Managing this threat requires means with which to convey the magnitude of the problem, forecast long-term trends, and assess the risks associated with changes in land use. Here we focus on approaches for assessing the risks of changes in land use, which include combining biological effect studies with catchment and hydrodynamic modeling, using statistical models that forecast the distribution and abundance of species relative to changes in habitat type, and using sensitive species that play important ecological roles as indicators of change.

Forecasting the limits of resilience: integrating empirical research with theory

Despite the increasing evidence of drastic and profound changes in many ecosystems, often referred to as regime shifts, we have little ability to understand the processes that provide insurance against such change (resilience). Modelling studies have suggested that increased variance may foreshadow a regime shift, but this requires long-term data and knowledge of the functional links between key processes. Field-based research and ground-truthing is an essential part of the heuristic that marries theoretical and empirical research, but experimental studies of resilience are lagging behind theory, management and policy requirements. Empirically, ecological resilience must be understood in terms of community dynamics and the potential for small shifts in environmental forcing to break the feedbacks that support resilience. Here, we integrate recent theory and empirical data to identify ways we might define and understand potential thresholds in the resilience of nature, and thus the potential for regime shifts, by focusing on the roles of strong and weak interactions, linkages in meta-communities, and positive feedbacks between these and environmental drivers. The challenge to theoretical and field ecologists is to make the shift from hindsight to a more predictive science that is able to assist in the implementation of ecosystem-based management.

Determining effects of suspended sediment on condition of a suspension feeding bivalve (Atrina zelandica): results of a survey, a laboratory experiment and a field transplant experiment

The horse mussel Atrina zelandica (Gray) is a large, suspension feeding pinnid bivalve, common in coastal and estuarine areas of northern New Zealand. As a suspension feeder, Atrina is likely to be influenced by suspended sediment loads. We conducted a laboratory experiment to determine the effect of short-term elevations in turbidity levels, such as those commonly recorded during storms, on the physiological condition and clearance rates of Atrina. We also conducted a field survey and a 3-month transplant experiment at multiple sites along a gradient of increasing suspended sediment load in a New Zealand estuary. Laboratory clearance rates of Atrina declined above a threshold suspended sediment concentration, and Atrina physiological condition at the end of this experiment was lower in high cf. low turbidity treatments. Decreases in Atrina condition were detected after exposure to elevated levels for only 3 days. The field survey and transplant experiment provided empirical evidence of a strong, negative effect of increasing suspended sediment flux on the physiological condition of Atrina. We suggest that relationships between the physiological condition of suspension feeders and sediment settling flux could provide a link between sediment inputs, which commonly occur as a result of catchment runoff during rainfall events, and the ecological health of estuarine and shallow coastal areas. Our study also demonstrated that Atrina have a natural distribution limit controlled by suspended sediment load. Thus, there is potential for larger-scale functional and structural effects on benthic communities in estuarine and coastal areas with high rates of sedimentation.

Functional Role of Large Organisms in Intertidal Communities: Community Effects and Ecosystem Function

In marine soft sediments, large organisms are potentially important players in the nonlinear interactions that occur among animals, their food, and their chemical environment, all of which influence the contribution of benthos to ecosystem function. We investigated the consequences of removing large individuals of two functionally contrasting benthic communities on nutrient regeneration, microphyte standing stock, and macrobenthic community composition. The experiment was conducted at two adjacent sites that were physically similar but biologically different, one dominated by large deposit feeders and the other by large suspension feeders. Chemical fluxes were measured in experimental plots, and sediments were sampled to assess changes in macrofauna, sediment grain size, organic content, and microphyte standing stock. Our results demonstrate that the removal of large suspension feeders or deposit feeders influenced the flux of nitrogen and oxygen, surficial sediment characteristics, and community composition. In the deposit-feeder community, interactions between nutrient regeneration and grazing highlight important feedbacks between large macrofauna and biogeochemical processes and production by microphytes, indicating that the loss of large infauna driven by increased rates of anthropogenic disturbance may lead to functional extinction and cause shifts in community structure and ecosystem performance.

Spatial structure of bivalves in a sandflat:: Scale and generating processes

A survey was conducted during the summer of 1994 within a fairly homogeneous 12.5 ha area of sandflat off Wiroa Island, in Manukau Harbour, New Zealand, to identify factors controlling the spatial distributions of the two dominant bivalves, Macomona liliana Iredale and Austrovenus stutchburyi (Gray), and to look for evidence of adult–juvenile interactions within and between species. Most of the large–scale spatial structure detected in the bivalve count variables (two species, several size classes of each) was explained by the physical and biological variables. The results of principal component analysis and spatial regression modelling suggest that different factors are controlling the spatial distributions of adults and juveniles. Larger size classes of both species displayed significant spatial structure, with physical variables explaining some but not all of this variation. Smaller organisms were less strongly spatially structured, with virtually all of the structure explained by physical variables. The physical variables important in the regression models differed among size classes of a species and between species. Extreme size classes (largest and smallest) were best explained by the models; physical variables explained from 10% to about 70% of the variation across the study site. Significant residual spatial variability was detected in the larger bivalves at the scale of the study site. The unexplained variability (20 to 90%) found in the models is likely to correspond to phenomena operating at smaller scales. Finally, we found no support for adult–juvenile interactions at the scale of our study site, given our sampling scale, after controlling for the effects of the available physical variables. This is in contrast to significant adult–juvenile interactions found in smaller–scale surveys and in field experiments. Our perception of adult–juvenile interactions thus depends on the scale of study.

THE EFFECTS OF HABITAT LOSS, FRAGMENTATION, AND COMMUNITY HOMOGENIZATION ON RESILIENCE IN ESTUARIES

When changes in the frequency and extent of disturbance outstrip the recovery potential of resident communities, the selective removal of species contributes to habitat loss and fragmentation across landscapes. The degree to which habitat change is likely to influence community resilience will depend on metacommunity structure and connectivity. Thus ecological connectivity is central to understanding the potential for cumulative effects to impact upon diversity. The importance of these issues to coastal marine communities, where the prevailing concept of open communities composed of highly dispersive species is being challenged, indicates that these systems may be more sensitive to cumulative impacts than previously thought. We conducted a disturbance-recovery experiment across gradients of community type and environmental conditions to assess the roles of ecological connectivity and regional variations in community structure on the recovery of species richness, total abundance, and community composition in Mahurangi Harbour, New Zealand. After 394 days, significant differences in recovery between sites were apparent. Statistical models explaining a high proportion of the variability (R2 > 0.92) suggested that community recovery rates were controlled by a combination of physical and ecological features operating across spatial scales, affecting successional processes. The dynamic and complex interplay of ecological and environmental processes we observed driving patch recovery across the estuarine landscape are integral to recovery from disturbances in heterogeneous environments. This link between succession/recovery, disturbance, and heterogeneity confirms the utility of disturbance-recovery experiments as assays for cumulative change due to fragmentation and habitat change in estuaries.

The Effect of Spatial and Temporal Heterogeneity on the Design and Analysis of Empirical Studies of Scale-Dependent Systems

Processes interacting across scales of space and time influence emergent patterns in ecological systems, but to obtain strong inference and empirical generalities, ecologists need to balance reality with the practicalities of design and analyses. This article discusses heterogeneity, scaling, and design analysis problems and offers potential solutions to improve empirically based research. In particular, we recommend bridging the dichotomy between correlative and manipulative studies by nesting manipulative studies within a correlative framework. We suggest that building on variation, by designing studies to detect variability, rather than fighting it often leads to an increase in generality. We also emphasize the importance of natural history information for determining likely scales of spatial and temporal heterogeneity and the probable occurrence of feedback loops, indirect effects, and interacting processes. Finally, we integrate these concepts and suggest planned iterations between multiscale studies to build up natural history information and test the strength of relationships across space and time. This offers a way forward in terms of heuristically developing models and determining ecological generalities.

Ocean acidification at high latitudes: potential effects on functioning of the Antarctic bivalve Laternula elliptica.

Ocean acidification is a well recognised threat to marine ecosystems. High latitude regions are predicted to be particularly affected due to cold waters and naturally low carbonate saturation levels. This is of concern for organisms utilising calcium carbonate (CaCO3) to generate shells or skeletons. Studies of potential effects of future levels of pCO2 on high latitude calcifiers are at present limited, and there is little understanding of their potential to acclimate to these changes. We describe a laboratory experiment to compare physiological and metabolic responses of a key benthic bivalve, Laternula elliptica, at pCO2 levels of their natural environment (430 µatm, pH 7.99; based on field measurements) with those predicted for 2100 (735 µatm, pH 7.78) and glacial levels (187 µatm, pH 8.32). Adult L. elliptica basal metabolism (oxygen consumption rates) and heat shock protein HSP70 gene expression levels increased in response both to lowering and elevation of pH. Expression of chitin synthase (CHS), a key enzyme involved in synthesis of bivalve shells, was significantly up-regulated in individuals at pH 7.78, indicating L. elliptica were working harder to calcify in seawater undersaturated in aragonite (ΩAr = 0.71), the CaCO3 polymorph of which their shells are comprised. The different response variables were influenced by pH in differing ways, highlighting the importance of assessing a variety of factors to determine the likely impact of pH change. In combination, the results indicate a negative effect of ocean acidification on whole-organism functioning of L. elliptica over relatively short terms (weeks-months) that may be energetically difficult to maintain over longer time periods. Importantly, however, the observed changes in L. elliptica CHS gene expression provides evidence for biological control over the shell formation process, which may enable some degree of adaptation or acclimation to future ocean acidification scenarios.