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Whereas the encrusting species Pseudosuberites sp. North Tower is the most intensively sampled of the Hikurangi sites [8] , [12] , [15] , [17] and illustrates most of the seep-associated habitats and fauna characteristic of the region Figure 4. The first signs of seep-associated habitats at the periphery of the seep were scattered Calyptogena sp. Authigenic carbonates occurred more frequently approaching the outskirts of the high backscatter SSS region and were often associated with low densities of Lamellibrachia sp.
At the boundary of the high-backscatter region, there was an abrupt transition from predominantly muddy sediments to authigenic carbonate blocks interspersed with dark sediments populated by ampharetids. Tubeworms were strongly associated with carbonates within the seep site Figure 5. Vesicomyid clam shells covered the seabed in localised patches m 2 scale but live Calyptogena sp. Transects are plotted against a side-scan sonar image of the seabed generated during SO darker pixels indicate stronger acoustic backscatter.
Data are from analyses of still images taken during towed camera transects across each site. Profiles are scaled to horizontal distance travelled x axis. South Tower. No additional biological sampling has been undertaken at South Tower since TAN, findings from which are described by Baco et al. Review of video and stills from the site in the present study confirmed habitats and fauna similar to those recorded at North Tower. Piwakawaka showed in SSS images as a region of stronger backscatter covering a seabed area of ca.
The site is similar to North Tower in terms of both substrata and fauna Figure 5 , but with low densities of bathymodiolin mussels in places. Tui forms the summit of a rise in the northern part of Opouawe Bank.
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The site was similar to North Tower in that the central area consisted of exposed authigenic carbonate blocks interspersed by muddy sediments, with tubeworms, clams, and ampharetid patches patchily distributed among them. It differed, however, in having a greater proportion of the substratum covered by Calyptogena sp.
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On muddy sediments surrounding the site, areas of disarticulated clam shells were common and often associated with populations of G. Takahe showed in SSS images as a roughly circular area of elevated backscatter ca. Patches of dark sediment with high densities of ampharetid polychaete pits and white bacterial mats were common Figure 5 , particularly on the north-eastern circumference of the site. Infrequent patches of Calyptogena sp. No other chemoautotrophic megafauna were observed.
On the last occasion this site was sampled SO, April , the dark, sulphide-rich sediments were less distinct in camera transects than in earlier years , Subsequent core samples showed that the sulphide-rich patches were overlain by deposits of fresh planktonic detrital matter, which obscured the dark sediments below. Hihi , LM , and Kereru. No additional sampling has been undertaken at these sites since TAN Habitats and fauna have been described by Baco et al.
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All three sites consist primarily of extensive weathered carbonate rock platforms with sparse Lamellibrachia sp. Scleractinian coral rubble and evidence of trawling abandoned gear and scour marks were widespread. Data, symbols, and scale as for Figure 5. Other than a single multicorer deployment at Bear's Paw during SO SO station , no additional biological sampling has been undertaken at these sites since SO Substratum and faunal characteristics have been described by Jones et al. Moa reef. As described by Jones et al. No live seep-associated fauna were recorded. Southwest Moa. Jones et al.
Additional SSS coverage collected during SO together with a single camera transect across the western extremity of the site confirmed that Southwest Moa is an area of active seepage supporting the highest densities of chemoautotrophic taxa recorded at any site. Figure 2 E , Figure 6. Stelletta n.
Rock Garden Knoll. Extensive weathered carbonate substratum was recorded but with no seep fauna recorded other than a single live Lamellibrachia sp. Faure Site. As described by Naudts et al. Still images were intermittent and of low quality at this site but video showed apparently live Bathymodiolin mussels associated with one of the areas of clam shell. Image quality here was also poor but video revealed extensive areas of weathered carbonate platform with widespread Calyptogena sp. No live seep fauna were recorded. As described by Naudts et al [34] , the only camera transect across this site showed extensive weathered carbonates with a single small patch ca.
Figure 6. Builder's Pencil is on a ridge of weathered carbonates and is remarkable for extensive areas of seabed overlain by accumulations of Calyptogena sp. Video analyses confirmed a seabed area of ca. Live Lamellibrachia sp. Bathymodiolin mussels occurred frequently but at low densities, and only one patch of live Calyptogena sp. On the flanks of the shell-covered ridge, carbonate rocks were colonised by sponges, crinoids, and antipatherian, stylasterid, and gorgonian corals.
Profiles across seep sites in each of the regions, using data from analysis of still images illustrate differences in substratum and community composition within and among regions Figures 5 and 6. Conspicuous distinctions among sites are the absence of carbonate rocks at Takahe, the extensive areas covered by Calyptogena sp.
The NMDS ordination based on observations of seep-associated fauna and corals in video transects Figure 7 indicated that communities range between three extreme states represented by transects at Takahe, Southwest Moa, and Moa Reef, respectively. The Takahe extreme was characterised primarily by the absence of all mega-epifauna other than highly localised populations of Calyptogena sp. The Southwest Moa extreme was characterised by the highest densities of all seep-associated taxa, and moderate areas of Calyptogena sp.
The third extreme, represented by Moa, was characterised by scleractinian corals and low densities, or absence of, chemoautotrophic taxa, including ampharetid patches and bacteria. Vectors of variable contributions Figure 7 B show that variability among sites along an axis from top left Takahe to bottom right Moa in the ordination was driven primarily by changes in the relative occurrence of bacterial mats and ampharetid patches increasing towards top left , and scleractinian corals increasing towards bottom right.
However, there was also marked variability along an axis orthogonal to this which was associated primarily with changes in the occurrences of live populations of the four megafaunal chemoautotrophic taxa; occurrence of clams, tube worms, mussels, and Stelletta n.
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The horizontal dimension of the ordination was correlated with the percentage of carbonate rock substratum Pearson correlation, 0. Underlying data are log-transformed occurrence records of chemoautotrophic megafauna Lamellibrachia sp. A — Transects distinguished by region symbols; see Figure 1 for context and seep site labels; see Table 1 for full names.
B — Contributions Pearson correlations of faunal variables to between-sample distances. The spread of data points between the three extreme states in the ordination indicated considerable variability at both within-region and within-site scales.
For instance, Opouawe Bank sites span a large proportion of the total ordination space, ranging from the extreme, no-carbonate, no-tubeworm, no-mussel state of Takahe to massive carbonates with large areas of Calyptogena sp. Similarly, in the Omakere Ridge region, Moa and Southwest Moa are adjoining sites yet contain strongly contrasting habitats and fauna; transects at these sites representing two of the three extreme states in the ordination. The ordination also indicated some separation between regions, particularly between Opouawe Bank and the other regions.
This was driven in part by higher incidence of bacterial mats and ampharetid patches at Opouawe Bank sites, but also by the presence of Stelletta n. However, there was convergence between sites independent of spatial separation. For instance, Tui and Builder's Pencil were similar, despite being located on Opouawe Bank and Ritchie Ridge, respectively; the similarity in this case being driven largely by high proportions of seabed covered by Calyptogena shells at both sites.
Comparing these values to the distribution of sites in the NMDS ordination Figure 7 C showed that increasing trawl intensity was associated with the gradient of decreasing occurrence of live chemoautotrophic fauna identified above, but was also inversely correlated with depth; shallower sites having fewer live fauna and greater trawl intensity.
Selected seep sites are identified by name see Table 1 for full names.