Seascape ecology of coastal biogenic habitats: advances, gaps, and challenges

We review the progress made in the emerging field of coastal seascape ecology, i.e. the application of landscape ecology concepts and techniques to the coastal marine environment. Since the early 1990s, the landscape ecology approach has been applied in several coastal subtidal and intertidal biogenic habitats across a range of spatial scales. Emerging evidence indicates that animals in these seascapes respond to the structure of patches and patch mosaics in different ways and at different spatial scales, yet we still know very little about the ecological significance of these relationships and the consequences of change in seascape patterning for ecosystem functioning and overall biodiversity. Ecological interactions that occur within patches and among different types of patches (or seascapes) are likely to be critically important in maintaining primary and secondary production, trophic transfer, biodiversity, coastal protection, and supporting a wealth of ecosystem goods and services. We review faunal responses to patch and seascape structure, including effects of fragmentation on 5 focal habitats: seagrass meadows, salt marshes, coral reefs, mangrove forests, and oyster reefs. Extrapolating and generalizing spatial relationships between ecological patterns and processes across scales remains a significant challenge, and we show that there are major gaps in our understanding of these relationships. Filling these gaps will be crucial for managing and responding to an inevitably changing coastal environment. We show that critical ecological thresholds exist in the structural patterning of biogenic ecosystems that, when exceeded, cause abrupt shifts in the distribution and abundance of organisms. A better understanding of faunal–seascape relationships, including the identifications of threshold effects, is urgently needed to support the development of more effective and holistic management actions in restoration, site prioritization, and forecasting the impacts of environmental change.

Patterns of scale-dependency and the influence of map resolution on the seascape ecology of reef fish

Detection and perception of ecological relationships between biota and their surrounding habitats is sensitive to analysis scale and resolution of habitat data. We measured strength of univariate linear correlations between reef fish and seascape variables at multiple spatial scales (25 to 800 m). Correlation strength was used to identify the scale that best associates fish to their surrounding habitat. To evaluate the influence of map resolution, seascape variables were calculated based on 4 separate benthic maps produced using 2 levels of spatial and thematic resolution, respectively. Individual seascape variables explained only 25% of the variability in fish distributions. Length of reef edge was correlated with more aspects of the fish assemblage than other features. Area of seagrass and bare sand correlated with distribution of many fish, not just obligate users. No fish variables correlated with habitat diversity. Individual fish species achieved a wider range of correlations than mobility guilds or the entire fish assemblage. Scales of peak correlation were the same for juveniles and adults in a majority of comparisons. Highly mobile species exhibited broader scales of peak correlation than either resident or moderately mobile fish. Use of different input maps changed perception of the strength and even the scale of peak correlations for many comparisons involving hard bottom edge length and area of sand, whereas results were consistent regardless of map type for comparisons involving area of seagrass and habitat diversity.

Practicing coastal seascape ecology

Landscape ecology concepts developed from terrestrial systems have recently emerged as theoretical and analytical frameworks that are equally useful for evaluating the ecological consequences of spatial patterns and structural changes in the submerged landscapes of coastal ecosystems. The benefits of applying a spatially-explicit perspective to resource management and restoration planning in the coastal zone are rapidly becoming apparent. This Theme Section on the application of landscape ecology to the estuarine and coastal environment emerged from a special symposium at the Coastal and Estuarine Research Federation (CERF) 20th Biennial Conference (Estuaries and Coasts in a Changing World) held in Portland, Oregon, USA, in November 2009. The 7 contributions in this Theme Section collectively provide substantial insights into the current status and application of the landscape approach in shallow marine environments, and identify significant knowledge gaps, as well as potential directions for the future advancement of ‘seascape ecology’.

Quantifying seascape structure: extending terrestrial spatial pattern metrics to the marine realm

Spatial pattern metrics have routinely been applied to characterize and quantify structural features of terrestrial landscapes and have demonstrated great utility in landscape ecology and conservation planning. The important role of spatial structure in ecology and management is now commonly recognized, and recent advances in marine remote sensing technology have facilitated the application of spatial pattern metrics to the marine environment. However, it is not yet clear whether concepts, metrics, and statistical techniques developed for terrestrial ecosystems are relevant for marine species and seascapes. To address this gap in our knowledge, we reviewed, synthesized, and evaluated the utility and application of spatial pattern metrics in the marine science literature over the past 30 yr (1980 to 2010). In total, 23 studies characterized seascape structure, of which 17 quantified spatial patterns using a 2-dimensional patch-mosaic model and 5 used a continuously varying 3-dimensional surface model. Most seascape studies followed terrestrial-based studies in their search for ecological patterns and applied or modified existing metrics. Only 1 truly unique metric was found (hydrodynamic aperture applied to Pacific atolls). While there are still relatively few studies using spatial pattern metrics in the marine environment, they have suffered from similar misuse as reported for terrestrial studies, such as the lack of a priori considerations or the problem of collinearity between metrics. Spatial pattern metrics offer great potential for ecological research and environmental management in marine systems, and future studies should focus on (1) the dynamic boundary between the land and sea; (2) quantifying 3-dimensional spatial patterns; and (3) assessing and monitoring seascape change.

Influence of seagrass landscape structure on the juvenile blue crab habitat-survival function

Organismal survival in marine habitats is often positively correlated with habitat structural complexity at local (within-patch) spatial scales. Far less is known, however, about how marine habitat structure at the landscape scale influences predation and other ecological processes, and in particular, how these processes are dictated by the interactive effect of habitat structure at local and landscape scales. The relationship between survival and habitat structure can be modeled with the habitat-survival function (HSF), which often takes on linear, hyperbolic, or sigmoid forms. We used tethering experiments to determine how seagrass landscape structure influenced the HSF for juvenile blue crabs Callinectes sapidus Rathbun in Back Sound, North Carolina, USA. Crabs were tethered in artificial seagrass plots of 7 different shoot densities embedded within small (1 – 3 m2) or large (>100 m2) seagrass patches (October 1999), and within 10 × 10 m landscapes containing patchy (<50% cover) or continuous (>90% cover) seagrass (July 2000). Overall, crab survival was higher in small than in large patches, and was higher in patchy than in continuous seagrass. The HSF was hyperbolic in large patches and in continuous seagrass, indicating that at low levels of habitat structure, relatively small increases in structure resulted in substantial increases in juvenile blue crab survival. However, the HSF was linear in small seagrass patches in 1999 and was parabolic in patchy seagrass in 2000. A sigmoid HSF, in which a threshold level of seagrass structure is required for crab survival, was never observed. Patchy seagrass landscapes are valuable refuges for juvenile blue crabs, and the effects of seagrass structural complexity on crab survival can only be fully understood when habitat structure at larger scales is considered.

PICES Press, Vol. 19, No. 2, Summer 2011

•The 2011 Inter-sessional Science Board Meeting: A Note from Science Board Chairman (pp. 1-4) •Indicators for Status and Change within North Pacific Marine Ecosystems: A FUTURE Workshop (pp. 5-8) •PICES Calendar (p. 8) •2011 ESSAS Open Science Meeting (pp. 9-13) •The 5th Zooplankton Production Symposium (pp. 14-17) •Workshop on "Individual-Based Models of Zooplankton” (pp. 18-21) •New Book Release on the 100th Anniversary of the T/S Osharu Maru (p. 21) •Workshop on “Advances in Genomic and Molecular Studies of Zooplankton” (pp. 22-24) •Workshop on “Updates and Comparisons of Zooplankton Time Series” (pp. 25-27) •Workshop on “Impacts of Ocean Acidification on Zooplankton” (pp. 28-29) •Workshop on “Automated Visual Plankton Identification” (p. 30) •Professor Plum in the Dining Room with a Knife (p. 31) •PICES and ICES on the River Elbe (p. 32) •The State of the Western North Pacific in the Second Half of 2010 (pp. 33-34) •The Bering Sea: Current Status and Recent Events (pp. 35-37) •Northeast Pacific News (pp. 38-39) •PICES Advice on Marine Ecology at a Canadian Judicial Inquiry (p. 40)

Diel movements of fishes linked to benthic seascape structure in a Caribbean coral reef ecosystem

Many common fishes associated with Caribbean coral reef ecosystems use resources from more than 1 patch type during routine daily foraging activities. Few studies have provided direct evidence of connectivity across seascapes, and the importance of benthic seascape structure on movement behavior is poorly known. To address this knowledge gap, we coupled hydro-acoustic technology to track fish with seafloor mapping and pattern analysis techniques from landscape ecology to quantify seascape structure. Bluestriped grunts Haemulon sciurus and schoolmaster snapper Lutjanus apodus were tracked over 24 h periods using boat-based acoustic telemetry. Movement pathways, and day and night activity spaces were mapped using geographical information system (GIS) tools, and seafloor structure within activity spaces was mapped from high-resolution aerial photography and quantified using spatial pattern metrics. For both fish species, night activity spaces were significantly larger than day activity spaces. Fish exhibited a daytime preference for seascapes with aggregate coral reef and colonized bedrock, then shifted to night activity spaces with lower complexity soft sediment including sand, seagrass, and scattered coral/rock. Movement path complexity was negatively correlated with seascape complexity. This demonstrates direct connectivity across multiple patch types and represents the first study to apply quantitative landscape ecology techniques to examine the movement ecology of marine fish. The spatially explicit approach facilitates understanding to the linkages between biological processes and the heterogeneity of the landscape. Such studies are essential for identifying ecologically relevant spatial scales, delineating essential fish habitat and designing marine protected areas.

Brevetoxin in two planktivorous fishes after exposure to Karenia brevis: implications for food-web transfer to bottlenose dolphins

Brevetoxin uptake was analyzed in 2 common planktivorous fish that are likely foodweb vectors for dolphin mortality events associated with brevetoxin-producing red tides. Fish were exposed to brevetoxin-producing Karenia brevis for 10 h under conditions previously reported to produce optimal uptake of toxin in blood after oral exposure. Striped mullet Mugil cephalus were exposed to a low dose of brevetoxin, and uptake and depuration by specific organs were evaluated over a 2 mo period. Atlantic menhaden Brevoortia tyrannus specimens were used to characterize a higher brevetoxin dose uptake into whole body components and evaluate depuration over 1 mo. We found a high uptake of toxin by menhaden, with a body to water ratio of 57 after a 10 h exposure and a slow elimination with a half life (t1/2) of 24 d. Elimination occurred rapidly from the intestine (t1/2 < 1 wk) and muscle (t1/2 ≈ 1 wk) compartments and redistributed to liver which continued to accumulate body stores of toxin for 4 wk. The accumulation and elimination characteristics of the vectoring capacity of these 2 fish species are interpreted in relation to data from the Florida Panhandle dolphin mortality event of 2004. We show that due to slow elimination rate of brevetoxin in planktivorous fish, brevetoxin-related dolphin mortality events may occur without evidence of a concurrent harmful algal bloom event.

Domoic acid contamination within eight representative species from the benthic food web of Monterey Bay, California, USA

Benthic food webs often derive a significant fraction of their nutrient inputs from phytoplankton in the overlying waters. If the phytoplankton include harmful algal species like Pseudo-nitzschia australis, a diatom capable of producing the neurotoxin domoic acid (DA), the benthic food web can become a depository for phycotoxins. We tested the general hypothesis that DA contaminates benthic organisms during local blooms of P. australis, a widespread toxin producer along the US west coast. To test for trophic transfer and uptake of DA into the benthic food web, we sampled 8 benthic species comprising 4 feeding groups: filter feeders (Emerita analoga and Urechis caupo); a predator (Citharichthys sordidus); scavengers (Nassarius fossatus and Pagurus samuelis) and deposit feeders (Neotrypaea californiensis, Dendraster excentricus and Olivella biplicata). Sampling occurred before, during and after blooms of P. australis in Monterey Bay, CA, USA during 2000 and 2001. DA was detected in all 8 species, with contamination persisting over variable time scales. Maximum DA levels in N. fossatus (674 ppm), E. analoga (278 ppm), C. sordidus (515 ppm), N. californiensis (145 ppm), P. samuelis (56 ppm), D. excentricus (15 ppm) and O. biplicata (3 ppm) coincided with P. australis blooms, while DA levels in U. caupo remained above 200 ppm (max. = 751 ppm) throughout the study period. DA in 6 species exceeded levels thought to be safe for higher level consumers (i.e. ≥20 ppm) and thus is likely to have deleterious effects on marine birds, sea lions and the endangered California sea otter, known to prey upon these benthic species.

Anti-grazing properties of the toxic dinoflagellate Karlodinium veneficum during predator-prey interactions with the copepod Acartia tonsa

Karlodinium veneficum (syn. Karlodinium micrum, Bergholtz et al. 2006; J Phycol 42:170–193) is a small athecate dinoflagellate commonly present in low levels in temperate, coastal waters. Occasionally, K. veneficum forms ichthyotoxic blooms due to the presence of cytotoxic, hemolytic compounds, putatively named karlotoxins. To evaluate the anti-grazing properties of these karlotoxins, we conducted food removal experiments using the cosmopolitan copepod grazer Acartia tonsa. Wild-caught, adult female A. tonsa were exposed to 6 monoalgal or mixed algal diets made using bloom concentrations of toxic (CCMP 2064) and non-toxic (CSIC1) strains of K. veneficum. Ingestion and clearance rates were calculated using the equations of Frost (1972). Exposure to the toxic strain of K. veneficum did not contribute to an increased mortality of the copepods and no significant differences in copepod mortality were found among the experimental diets. However, A. tonsa had significantly greater clearance and ingestion rates when exposed to a monoalgal diet of the non-toxic strain CSIC1 than when exposed to the monoalgal diet of toxic strain CCMP 2064 and mixed diets dominated by this toxic strain. These results support the hypothesis that karlotoxins in certain strains of K. veneficum deter grazing by potential predators and contribute to the formation and continuation of blooms.

Prevalence of brevetoxins in prey fish of bottlenose dolphins in Sarasota Bay, Florida

Blooms of the brevetoxin-producing dinoflagellate Karenia brevis have been linked to high mortality of bottlenose dolphins Tursiops truncatus on Florida’s Gulf of Mexico coast. A clear understanding of trophic transfer of brevetoxin from its algal source up the food web to top predators is needed to assess exposure of affected dolphin populations. Prey fish constitute a means of accumulating and transferring brevetoxins and are potential vectors of brevetoxin to dolphins frequently exposed to K. brevis blooms. Here we report results of brevetoxin analyses of the primary fish species consumed by long-term resident bottlenose dolphins inhabiting Sarasota Bay, Florida. Fish collected during K. brevis blooms in 2003 to 2006 were analyzed by competitive enzyme-linked immunosorbent assay (ELISA) and had brevetoxin concentrations ranging from 4 to 10844 ng PbTx-3 eq g–1 tissue. Receptor binding assay (RBA) and liquid chromatography–mass spectrometry (LC-MS) analysis confirmed toxicity and the presence of parent brevetoxins and known metabolites. Fish collected in the absence of K. brevis blooms tested positive for brevetoxin by ELISA and RBA, with concentrations up to 1500 ng PbTx-3 eq g–1 tissue. These findings implicate prey fish exposed to K. brevis blooms as brevetoxin vectors for their dolphin predators and provide a critical analysis of persistent brevetoxin loads in the food web of dolphins repeatedly exposed to Florida red tides.

A case study on the impact of industrial effluent disposal on the fishery of Amba River estuary, Maharashtra

The impact of waste discharge on fishery resources is a matter of great concern. The accepted norm in all environmental impact assessment studies is to avoid areas of high fishery potential while locating a marine outfall. Contemplating on this aspect a case study was conducted in the Amba River estuary before and after the establishment of a petrochemical complex at Nagothane. The treated wastewater from this complex is released through a subsurface outfall after adopting effective control measures for marine disposal of waste. Experimental trawling was done at five locations covering a distance of 30 km during 1990 to 1991. The catch rate within the estuary varied from 0.6 to 255 kg/h (av 24 kg/h). The trend indicated considerable decrease in fishery potential from the mouth of the estuary (av 64 kg/h) to the upstream location (av 11 kg/h). A total of 49 species of fishes, 16 species of prawns, 7 species of crabs and 1 species of lobster were identified from the collections. Number of species gradually increased from the interior segment at Dharamtar (8) to the outer area near Revas (18). A comparison of the quantitative and qualitative nature of the post outfall and pre outfall data revealed only marginal difference. The study indicates that if necessary precautions are taken to render the waste harmless the marine ecology will hardly be affected.