Complex biogenic habitats formed by corals are important components of the megabenthos of seamounts, but their fragility makes them susceptible to damage by bottom trawling. Here we examine changes to stony corals and associated megabenthic assemblages on seamounts off Tasmania (Australia) with different histories of bottom-contact trawling by analysing 64 504 video frames (25 seamounts) and 704 high-resolution images (7 seamounts). Trawling had a dramatic impact on the seamount benthos: (1) bottom cover of the matrix-forming stony coral Solenosmilia variabilis was reduced by 2 orders of magnitude; (2) loss of coral habitat translated into 3-fold declines in richness, diversity and density of other megabenthos; and (3) megabenthos assemblage structures diverged widely between trawled and untrawled seamounts. On seamounts where trawling had been reduced to < 5% a decade ago and ceased completely 5 yr ago, there was no clear signal of recovery of the megabenthos; communities remained impoverished comprising fewer species at reduced densities. Differences in community structure in the trawled (as compared to the untrawled) seamounts were attributed to resistant species that survived initial impacts, others protected in natural refugia and early colonisers. Long-term persistence of trawling impacts on deep-water corals is consistent with their biological traits (e.g. slow growth rates, fragility) that make them particularly vulnerable. Because recovery on seamounts will be slow, the benefits from fishery closures may not be immediately recognisable or measureable. Spatial closures are crucial conservation instruments, but will require long-term commitments and expectations of performance whose time frames match the biological tempo in the deep sea.
Deep-sea fisheries operate globally throughout the world's oceans, chiefly targeting stocks on the upper and mid-continental slope and offshore seamounts. Major commercial fisheries occur, or have occurred, for species such as orange roughy, oreos, cardinalfish, grenadiers and alfonsino. Few deep fisheries have, however, been sustainable, with most deep-sea stocks having undergone rapid and substantial declines. Fishing in the deep sea not only harvests target species but can also cause unintended environmental harm, mostly from operating heavy bottom trawls and, to a lesser extent, bottom longlines. Bottom trawling over hard seabed (common on seamounts) routinely removes most of the benthic fauna, resulting in declines in faunal biodiversity, cover and abundance. Functionally, these impacts translate into loss of biogenic habitat from potentially large areas. Recent studies on longline fisheries show that their impact is much less than from trawl gear, but can still be significant. Benthic taxa, especially the dominant mega-faunal components of deep-sea systems such as corals and sponges, can be highly vulnerable to fishing impacts. Some taxa have natural resilience due to their size, shape, and structure, and some can survive in natural refuges inaccessible to trawls. However, many deep-sea invertebrates are exceptionally long-lived and grow extremely slowly: these biological attributes mean that the recovery capacity of the benthos is highly limited and prolonged, predicted to take decades to centuries after fishing has ceased. The low tolerance and protracted recovery of many deep-sea benthic communities has implications for managing environmental performance of deep-sea fisheries, including that (i) expectations for recovery and restoration of impacted areas may be unrealistic in acceptable time frames, (ii) the high vulnerability of deep-sea fauna makes spatial management—that includes strong and consistent conservation closures—an important priority, and (iii) biodiversity conservation should be > balanced with options for open areas that support sustainable fisheries.
SignificanceWe conducted a systematic, high-resolution analysis of bottom trawl fishing footprints for 24 regions on continental shelves and slopes of five continents and New Zealand. The proportion of seabed trawled varied >200-fold among regions (from 0.4 to 80.7% of area to a depth of 1,000 m). Within 18 regions, more than two-thirds of seabed area remained untrawled during study periods of 2–6 years. Relationships between metrics of total trawling activity and footprint were strong and positive, providing a method to estimate trawling footprints for regions where high-resolution data are not available. Trawling footprints were generally smaller in regions where fisheries met targets for exploitation rates, implying collateral environmental benefits of effective fisheries management.
Because the nature, tempo and trajectories of biological changes that follow the cessation of trawling are unknown for seamounts, it is unclear whether closing them to trawling will lead to a recovery of the fauna and, if so, over what time scales. This paper reports on a 'test of recovery' from repeated towed camera surveys on three seamounts off New Zealand in 2001 and 2006 (5 years apart) and three off Australia in 1997 and 2006 (10 years apart). In each region, seamounts where trawling had ceased were compared to adjacent seamounts where trawling was still active, and to seamounts that had never been trawled. If recovery signals existed, the likelihood of detecting them was high because the seamounts were relatively small and topographically simple, and because quantitative survey methods were employed. Multivariate patterns showed no change in the megafaunal assemblage consistent with recovery over a 5-10 year timeframe on seamounts where trawling had ceased. Results based on the number of species and diversity were equivocal, with some cases of increase and decrease on seamounts where trawling had ceased. A few individual taxa were found at significantly higher abundance in the later surveys where trawling had occurred. We suggest this may have resulted from their resistance to the direct impacts of trawling (two chrysogorgid corals and solitary scleractinians), or from protection in natural refuges inaccessible to trawls (unstalked crinoids, two chrysogorgid corals, gorgonians, and urchins). Alternatively, these taxa may represent the earliest stages of seamount recolonisation. They have potential to be dominant for long periods because the pre-trawling composition of benthic assemblages on seamounts includes taxa that grow slowly and ⁄ or have an association with 'thickets' of a single keystone stony coral (Solenosmilia variabilis) that has generated biogenic habitat over millennia. Resilience of seamount ecosystems dominated by corals is low compared to most other marine systems subject to disturbance by bottom trawling because there are no alternative habitats of the same value for supporting associated species, and because trawling typically removes coral habitat from large areas of individual seamounts. Management to conserve seamount ecosystems needs to account for changing oceanographic conditions (ocean acidification), as well as the direct impacts of human activities such as bottom trawling. Networks of spatial closures that include intact habitats over a range of depths, especially <1500 m, and on clusters and isolated seamounts, may be effective by maintaining the resilience of seamount benthic communities.
Imagery collected by still and video cameras is an increasingly important tool for minimal impact, repeatable observations in the marine environment. Data generated from imagery includes identification, annotation and quantification of biological subjects and environmental features within an image. To be long-lived and useful beyond their project-specific initial purpose, and to maximize their utility across studies and disciplines, marine imagery data should use a standardised vocabulary of defined terms. This would enable the compilation of regional, national and/or global data sets from multiple sources, contributing to broad-scale management studies and development of automated annotation algorithms. The classification scheme developed under the Collaborative and Automated Tools for Analysis of Marine Imagery (CATAMI) project provides such a vocabulary. The CATAMI classification scheme introduces Australian-wide acknowledged, standardised terminology for annotating benthic substrates and biota in marine imagery. It combines coarse-level taxonomy and morphology, and is a flexible, hierarchical classification that bridges the gap between habitat/biotope characterisation and taxonomy, acknowledging limitations when describing biological taxa through imagery. It is fully described, documented, and maintained through curated online databases, and can be applied across benthic image collection methods, annotation platforms and scoring methods. Following release in 2013, the CATAMI classification scheme was taken up by a wide variety of users, including government, academia and industry. This rapid acceptance highlights the scheme’s utility and the potential to facilitate broad-scale multidisciplinary studies of marine ecosystems when applied globally. Here we present the CATAMI classification scheme, describe its conception and features, and discuss its utility and the opportunities as well as challenges arising from its use.
Seamounts have often been viewed as specialized habitats that support unique communities; this notion has given rise to several hypotheses about how seamount ecosystems are structured. One, the ‘seamount oasis hypothesis’, predicts that invertebrates are more abundant, speciose and attain higher standing stocks on seamounts compared to other deep‐sea habitats. Because this hypothesis has remained untested for biomass, we ask two questions: (i) Do seamounts support a higher benthic biomass than nearby slopes at corresponding depths? (ii) If they do, which particular taxa and trophic groups drive observed difference in biomass? Analysis of more than 5000 sea‐floor images reveals that the mean biomass of epibenthic megafauna on 20 southwest Pacific seamounts was nearly four times greater than on the adjacent continental slope at comparable depths. This difference is largely attributable to the scleractinian coral Solenosmilia variabilis, whose mean biomass was 29 times higher on seamounts. In terms of trophic guilds, filter‐feeders and filter‐feeders/predators made up a significantly greater proportion of biomass on seamounts, whereas deposit feeders and those with mixed feeding modes dominated at slope habitats. Notwithstanding support for the seamount oasis hypothesis provided by this study, the hypothesis needs to be critically tested for seamounts in less productive regions, for seamounts with a greater proportion of soft substratum, and in other parts of the oceans where scleractinian corals are not prevalent. In this context, testing of seamount paradigms should be embedded in a broader ecological context that includes other margin habitats (e.g. canyons) and community metrics (e.g. diversity and body size).
The first large systematic collection of benthic invertebrate megafauna from the Australian continental margin (depths > 100 m) revealed high species richness and novelty on the south-western continental slope (100-1100 m depth; 18°S-35°S). A total of 1979 morphologically defined species was discriminated in seven taxa across all samples: Demospongiae, Decapoda, corals (Octocorallia and Antipatharia), Mollusca, Echinodermata, Ascidiacea, and Pycnogonida. Collectively, 59% were estimated to be new or unnamed species. The distribution pattern of megafaunal communities, analysed with a suite of 17 physical covariates, was most influenced at large spatial scales (>100s km) by bottom temperature, oxygen concentration and latitude, whereas at smaller scales (10s of km), seabed type was most influential. Many covariates are driven by the same physical processes and are correlated (e.g. to depth or latitude), thus it is not possible to ascribe causal relationships to fauna distributions. However, their identification highlights the spatial scales that determine the composition of megafaunal communities. Regional-scale transitions in bottom temperature and oxygen concentration are determined by water masses and currents that interact with the south-western margin seabed in different ways depending on location. The nested, smaller-scale heterogeneity of seabed type, classified simply as 'hard' or 'soft' terrain, differentiates consolidated attachment sites for sessile fauna from sediments suited to mobile and burrowing fauna. Different physical factors affect the distribution of benthic fauna at different scales. Collectively, these patterns of heterogeneity can be represented in a hierarchical framework that consists of biogeographic provinces, biomes, biogeomorphic features, terrains, and finer scales. The Australian government is using a hierarchical approach to identify bioregions for management purposes; a key aim is to ensure that a National Representative System of Marine Protected Areas (NRSMPA) will meet the requirement of comprehensiveness, adequacy and representativeness. Important findings from this study are that the provincial structure of invertebrate megabenthos broadly aligns with the provincial structure derived earlier from the distribution of fishes, but there are differences in the distribution of individual major taxa at both provincial and megahabitat scales. Representative coverage of rarer taxa or narrowly distributed taxa might not be feasible at the same time as ensuring main fauna groups are adequately represented. The hierarchical scales of heterogeneity of the megabenthos in this area, the differences between taxa, and the high proportion of apparently rare species make it clear that it will be as important to manage the area outside the NRSMPA as to manage the NRSMPA itself. Management will be required at different scales that correspond to the multiscale spatial heterogeneity of continental margin fauna.
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