Rapid assembly of Invertebrate data for the SE Regional Marine Plan. T.D. O’Hara, G.C.B. Poore, S. Ahyong, D.A. Staples Museum Victoria September 2002 2 Table of Contents Abstract ...................................................................................................................... 6 1. Introduction............................................................................................................ 8 The tasks for this project included ......................................................................... 8 2. SE continental slope .............................................................................................. 9 2.1. Methods........................................................................................................... 9 2.2. Results ........................................................................................................... 12 2.3. Summary ....................................................................................................... 13 2.4. East vs west fauna ......................................................................................... 13 2.5. North vs South Fauna.................................................................................... 14 2.6. Eastern Tasman Rise ..................................................................................... 14 2.7. Comparisons between taxonomic groups ..................................................... 14 2.8. Change in distribution with depth ................................................................. 16 2.9. Species endemic to the SE region ................................................................. 17 3. Macquarie Ridge .................................................................................................. 19 3.1. Methods......................................................................................................... 19 3.2. Results ........................................................................................................... 21 3.3. Summary ....................................................................................................... 21 4. Conclusions .......................................................................................................... 22 4. Participants .......................................................................................................... 23 5. References ............................................................................................................ 23 3 List of tables and figures Figure 1. Map showing the SE region and some of the place names used in the text. .. 7 Table 1. Geographic spread of the number of species recorded for each 0.5 degree of latitude (including interpolated records). ............................................................. 10 Table 2. Geographic spread of the number of collection stations by latitude (rows) and longitude (columns). Heavily surveyed cells include the Tasmania seamounts (44ºS) and off far-eastern Victoria (37.5-38.0ºS, 149-150ºE). Few samples have been collected from the East (43.5ºS, 149-150ºE) or South Tasman (<44ºS) Rises. .................................................................................................................... 11 Table 4. Spearman Rank correlation coefficients between the entire dataset and selected taxonomic groups for each depth strata on either side of the SE region. The number of cells of latitude is kept constant even if not all taxonomic-groups were present. ........................................................................................................ 15 Table 5. Spearman Rank correlation coefficients between the entire dataset and selected taxonomic groups for each depth strata on either side of the SE region. Only the cells of latitude that contained all taxonomic groups are included in the analysis. ................................................................................................................ 16 Table 5. Apparently endemic species from the continental slope of the SE region. ... 18 Table 6. Geographic spread of the number of species recorded in each cell from the Macquarie Ridge (including interpolated records). ............................................. 20 Table 7. Geographic spread of the number of collection stations by latitude (rows) and longitude (columns) from the Macquarie Ridge. Heavily surveyed cells include the Macquarie Island (54.5ºS) cell. ...................................................................... 20 Figure 2. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 200-500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 24 Figure 3. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 500-1000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 25 Figure 4. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 1500-2000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 26 Figure 5. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 1500-2000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 27 Figure 6. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 2000-2500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 28 Figure 7. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from below 2500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 29 Figure 8. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the western SE Australian continental slope from 200-500 m. 4 Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 30 Figure 9. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the western SE Australian continental slope from 500-1000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. .......................................................................................... 31 Figure 10. Stacked plot of the number of species per grid. ......................................... 32 Figure 11. Stacked plot of the species richness for each 100 m depth strata for all stations across the SE region................................................................................ 33 Figure 12. Species richness from various collection methods for 100 m depth strata across the SE region. ............................................................................................ 34 Figure 13. Species richness, species turnover and effort (no of samples) for 100 m depth strata across the SE region. ........................................................................ 34 Figure 14. Cluster diagram of all samples from the SE region collected using an epibenthic sled (species abundance data is square root transformed). ................. 35 Figure 15. MDS ordinations of species from the Macquarie Ridge, based on the presence/absence of each species along the Ridge in cells of one-degree latitude. Macquarie Island is located at 54ºS. .................................................................... 36 Figure 16. Cluster diagram of species from the Macquarie Ridge, based on the presence/absence of each species along the Ridge in cells of one-degree latitude between 500-1000 m. Macquarie Island is located at 54ºS ................................. 37 5 Abstract The distribution of echinoderms, decapods and pycnogonids from 1) the continental slope (>200 m) of the Australian SE region, and 2) the continental shelf and slope around Macquarie Island are analysed for geographical and bathymetric patterns. The patterns along the SE continental slope are indicative of a gradient from a temperate fauna of the Tasman Sea to the subantarctic fauna of the South Tasman Rise. This temperate fauna persists as far as mid Tasmania (approximately 42ºS), and possibly will be found further south with increased collection effort. Subantarctic species become progressively more important in the region south of Tasmania. The fauna on the Southern Tasmania seamounts is particularly distinctive. There is a large apparent difference between the eastern and western slopes, and much of this is due to the greater collection effort off the East Coast. Nevertheless, there is a small fauna (1.6%) that is only known from the western slope, and a larger (but as yet unquantified) fauna that is restricted to the Tasman Sea. The low collection effort precludes an analysis of the fauna of the southern and eastern Tasman Rises. There is a rapid turnover of species occurrence between the shelf fauna and the upper slope (150-300 m), followed by gradual change until 2500 m, after which there is another pronounced faunal change. There are no samples from the abyssal plain (>4000 m). Communities were divided into shelf (<200 m), upper slope (200-1000 m) and mid-lower slope categories (>1000 m). For at least one taxonomic group (ophiuroids) there is a peak of diversity at mid-slope (1000 m). There is a gradient along the Macquarie Ridge also, with the fauna of the Northern and Southern Ridges being distinct from each other and the fauna around Macquarie Island. The Northern Ridge supports subantarctic New Zealand species and the Southern Ridge Antarctic species. The shallow water fauna around the island is composed of shelf species from New Zealand and subantarctic species that have presumably arrived on kelp holdfasts from islands further west. However, this study must be considered preliminary as uneven sampling effort, both geographically and bathymetrically, is a major impediment to fine-scale analysis. 6 Port Macdonnell Bass Strait Gabo Is East Tasman Rise Tasmanian Seamounts South Tasman Rise 147°E Macquarie Island (54S) Figure 1. Map showing the SE region and some of the place names used in the text. 7 So Ma 1. Introduction Museum Victoria, in conjunction with the Australian Museum and CSIRO, was contracted by the National Oceans Office to prepare information on the community composition and distribution of key invertebrate groups (echinoderms, decapods and pycnogonids) on the outer shelf and slope of three Large Marine Domains so that it is available to be integrated with other datasets to develop an Interim Bioregionalisation for the South-east Marine Region. The tasks for this project included A matrix analysis of community composition of the benthic fauna along the eastern continental slope from the northern limit of the SE region (southern NSW) to the Tasmanian seamounts. The presence of each echinoderm, decapod and pycnogonid species will be scored for cells of resolution 0.5 degrees latitude by 200 m depth. Gaps in sampling effort will be compensated by interpolating from neighbouring cells. The final matrix will be analysed using multivariate statistics. A comparison of the fauna from the continental slope on the eastern and western sides of Bass Strait. A comparison of the fauna from the north and south Macquarie Ridges. A comparison of the fauna from the SE Tasmanian slope and the Eastern Tasman Rise. The target taxa for this project were echinoderms, decapods and pycnogonids. The rationale is that for these taxa 1) many specimens had already been identified in various museums and 2) taxonomic experts were readily available. Thus both existing and new data could be assembled within the limited timeframe available for this task (3 months). Although sessile groups such as cnidarians, bryozoans and sponges would have been very useful, a comprehensive dataset could not have been accumulated in this timeframe. Identification of these groups would be a major taxonomic exercise requiring many months if not years of effort. Pycnogonids were selected as a target group as they have a close association with hydroids and bryozoans may reflect the same factors influencing sessile-faunal community structure. 8 2. SE continental slope 2.1. Methods Data Most material was collected by the Endeavour (1910-1913), Kimbla (1973), BSS (1979-1983), Franklin (1986-89), Soela (1982-88), Kapala (1973-97) and CSIRO Southern Surveyor (1997, 2000) research vessels. This was supplemented by material collected on an ad-hoc basis from Fisheries Vessels. In all there were 574 species in 5100 collection (species/station) lots representing 61670 individual specimens. The material was collected using bottom trawls, an epibenthic sled, box corer and traps, but predominantly by trawling. The material is stored at Museum Victoria (MV), the Australian Museum (AM), the Tasmanian Museum and Art Gallery (TM) and CSIRO Marine Laboratories. Material at the South Australian Museum was not accessed due to lack of time. The dataset for this analysis included Echinodermata (Ophiuroidea, Holothuroidea), Decapoda, and Pycnogonida collected below 200 m). Although identified, asteroids, echinoids and crinoids were excluded from the final analyses, as the resulting dataset was not geographically comprehensive enough for a balanced analysis. Analysis The data were summarised into the presence/absence of each species in cells of one half a degree latitude for each of the eastern and western continental margins of the region (36-47ºS). The amount of longitude covered in each grid varied with the width of the continental slope at each latitude. For both the western and eastern slope, two additional cells were created representing the northern (<=35ºS) and southern (>=48ºS) borders of the region based on whether each species is known to occur at more northern or southern latitudes. The data were sourced from published literature and collection databases. Although the target depth segment was 200 m, not enough material was present at all longitudes to facilitate stratification of the dataset in this way. Consequently, depth strata of 200-500, 500-1000, 1000-1500, 1500-2000, 2000-2500, 2500+ m were finally used in the analysis. A species that occurs at the limit between depth strata (eg 500 m) was included in the next depth strata (500-1000 m). The actual data were supplemented by interpolated data, generated by recording a species as present for cells that are between known occurrences for each depth strata. The data were analysed using multivariate statistics, including MDS ordinations and cluster diagrams, using Bray-Curtis similarity coefficients and group-average linkage. The use of the Bray-Curtis similarity measure on presence/absence data is equivalent to using the Sorensen Binary coefficient. Patterns generated by different taxonomic groups were compared to the full dataset by Spearman Rank correlating the underlying similarity matrices. 9 Table 1. Geographic spread of the number of species recorded for each 0.5 degree of latitude (including interpolated records). Eastern slope (>=147ºE) Depth (m) 35+ 36.0 36.5 37.0 37.5 38.0 38.5 39.0 39.5 40.0 40.5 41.0 41.5 42.0 42.5 43.0 43.5 44.0 44.5 45.0 45.5 46.0 46.5 47.0 47.5 48+ 200-500 35 41 44 56 69 85 52 54 51 43 41 39 41 28 30 5 3 2 2 2 2 2 2 2 2 6 500-1000 63 60 68 84 82 105 66 68 65 62 58 61 58 55 33 23 20 30 10 10 10 10 10 10 10 10 1000-1500 52 41 43 41 41 90 52 49 49 49 49 50 48 46 42 37 37 61 10 10 10 10 10 14 15 15 1500-2000 46 33 34 34 34 53 51 38 44 24 24 24 33 23 16 16 16 37 2 2 2 2 2 2 2 2 2000-2500 12 5 5 5 5 5 18 7 7 9 9 9 21 20 12 7 7 14 4 4 4 4 4 4 4 4 2500+ 7 2 3 2 2 2 11 5 5 5 10 5 17 16 4 4 4 2 0 0 0 0 0 0 0 0 Western slope (<147ºE) Depth (m) 35+ 36.0 36.5 37.0 37.5 38.0 38.5 39.0 39.5 40.0 40.5 41.0 41.5 42.0 42.5 43.0 43.5 44.0 44.5 45.0 45.5 46.0 46.5 47.0 47.5 48+ 200 22 15 15 20 14 14 24 18 12 16 12 17 5 1 1 3 0 0 0 0 0 0 0 0 0 6 500 13 8 9 15 14 14 34 17 15 16 17 20 16 10 9 9 10 28 4 4 4 4 4 4 4 10 1000 9 4 4 4 3 5 9 4 4 4 5 6 5 5 5 7 7 26 6 6 6 6 6 6 6 15 1500 5 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 4 0 0 0 2 2000 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 2500 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 Table 2. Geographic spread of the number of collection stations by latitude (rows) and longitude (columns). Heavily surveyed cells include the Tasmania seamounts (44ºS) and off far-eastern Victoria (37.5-38.0ºS, 149-150ºE). Few samples have been collected from the East (43.5ºS, 149-150ºE) or South Tasman (<44ºS) Rises. Latitude 36.0ºS 137ºE 138 139 140 141 142 143 144 145 146 147 148 149 150ºE 16 36.5ºS 1 37.0ºS 5 37.5ºS 11 4 5 1 18 36 63 6 38.0ºS 4 2 8 78 38.5ºS 1 22 8 9 39.0ºS 5 20 39.5ºS 1 10 5 40.0ºS 3 4 1 40.5ºS 11 1 1 41.0ºS 4 4 7 1 41.5ºS 6 13 1 42.0ºS 2 6 3 42.5ºS 23 43.0ºS 43.5ºS 7 4 2 1 44.0ºS 3 2 2 2 1 1 37 44.5ºS 45.0ºS 45.5ºS 46.0ºS 1 46.5ºS 47.0ºS 47.5ºS 3 4 1 11 2.2. Results Eastern slope Depth strata 200-500 m (Figure 2) Multivariate analysis There is no area on the South Tasman Rise (<44ºS) at this depth. The next shallow water fauna south of 48ºS is at Macquarie Island, which is composed predominantly of subantarctic species. Southern Tasmanian cells (43-43.5 ºS) are outliers due to lack of sampling. The rest of the cells form three clusters based on S. NSW, E Bass Strait, and E Tasmania. Breaks occur at 37ºS and 40ºS. There is a peak in collection effort off far eastern Vic. However, many of these records are rare species, which contribute little to the multivariate analysis. 500-1000 m (Figure 3) The South Tasman Rise (45-47ºS) is an outlier due to lack of sampling at this depth. The other cells form three clusters based on NSW, E Tas and S Tas, with breaks at 38ºS and 42ºS. 1000-1500 m (Figure 4) This depth strata is very similar to the 500-1000 m strata with breaks between the three main clusters near 38ºS and 42ºS. There is some differentiation between the southern and northern South Tasman Rise, although this is based on relatively sparse data. The intensive collecting effort at the Tasmanian seamounts (44ºS) and off Gabo Island (38ºS) can be observed. 1500-2000 m (Figure 5) Again very similar, except that the Gabo Island no longer represents a peak of collection effort. The Tasmanian seamounts are still distinct at 44ºS. The three main clusters again break near 38ºS and 42ºS 2000-2500 m (Figure 6) Collection effort at this depth is patchy, mainly confined to transects off Eastern Bass Strait (38.5, 40, 40.5ºS) and SE Tasmania (41.5, 42ºS). The cluster diagram shows a gradient from NSW south to the seamounts with the South Tasman Rise as an outlier. However, this pattern is liable to be an artefact of the collection effort. 2500+ m (Figure 7) Similar to 2000-2500 m. 12 Western slope Depth strata 200-500 m (Figure 8) Multivariate analysis The southern cells (41.5 to 43ºS) form an outlying cluster due to lack of sampling. The rest of the cells form two clusters based on SA to W. Bass Strait (36-39ºS), and the other NW Tasmania (39.5-40.5ºS). 500-1000 m (Figure 9) 1000-2500+ m There is only half the number of records of the 200-500 m strata with peaks of collection effort off Port Macdonnell/Portland (38.5ºS), and the seamounts (44ºS). The South Tasman Rise (44.5-47ºS) is an outlier cluster due to lack of sampling at this depth. The other cells form three clusters based on SA, E Tas and S Tas, with breaks at 38ºS and 42ºS. S. Tasmania clusters with the South Tasman Rise. Collection effort is not adequate to analyse the fauna at this depth. 2.3. Summary Eastern slope Excluding outliers, the dendrograms for shallow depths (200-500 m) formed three main clusters, based on southern NSW (36-37ºS), eastern Bass Strait (37.5-39.5ºS) and eastern Tasmania (40-42.5ºS). Unfortunately there were few records from this depth from off Southern Tasmania. Further down the slope (500-2000 m), excluding the South Tasman Rise, again there were three main clusters based around NSW (to 37.5ºS), eastern Bass Strait and eastern Tasmania (38-42ºS), and southern Tasmania, including the seamounts (42.5-44ºS). From 1000-2000 m the Tasmanian seamounts become distinct, partly because of collection effort, but also due to the unusual fauna that is present. The South Tasman Rise is distinct on most analyses. However, some of this effect was liable to be due to lack of collection effort (Table 1). Western slope Due to low collection effort only the upper slope (200-1000 m) contained a dataset that was worth analysing. There is a gradient in species composition from off SE South Australia to southern Tasmania, with breaks at 39ºS and 41.5ºS for 200-500 m, and 37ºS and 42ºS at 500-1000 m. However, Southern Tasmania may be a cluster due to a collection artefact, as there are relatively few stations from that area (Table 2). The Tasmanian seamounts (44ºS) are distinctive. 2.4. East vs west fauna Of the 328 species included in this analysis (ophiuroids, holothurians, decapods and pycnogonids only), 205 were only recorded from the eastern slope, 19 only from the western slope and 104 from both eastern and western slopes. However, the difference between east and west was at least partly due to the much greater collection effort on the eastern slope. Many of the eastern-only species may be found on the western slope 13 with additional collection effort. Nevertheless at least some of these species are likely to be eastern slope endemics. A distinct eastern coast fauna has been found for shallow water (< 100 m) decapods and echinoderms (O’Hara & Poore 2000). Of more interest were the 19 species only found on the western slope. Of these nine were collection artefacts of this study and have been found in other surveys off central eastern Australia, often in shallower water (<200 m). Eight others are undescribed and may have a wider distribution when they are better known. Only two species, the ophiuroids Ophiomusium australe and Ophiomusium anisacanthum appear to be confirmed western slope endemics, only 1.6 % of the western fauna. These three species are common on the western slope across into South Australia. 2.5. North vs South Fauna There was a decline in species richness per cell with increasing latitude (Figure 10) on both the eastern and western slopes. Again this may be largely due to the uneven sampling effort, although a latitudinal decline in species richness has been noted for many other habitats (O’Hara & Poore 2000). The slopes off eastern and western Bass Strait were the most heavily sampled areas and the areas with greatest species richness. Another heavily sampled area, the Tasmanian seamounts, also shows as a spike of diversity at 44ºS. The turnover of species from north to south was also evident from Figure 10. There was a widespread fauna, comprising 13% of the total fauna, which occurs all along the slope as far as the Tasmanian seamounts. There was a Northern fauna (species that do not occur >44ºS), which is numerically dominant (44 % of the total fauna) which persisted to 42.5ºS on the eastern side and 41.5ºS on the west. There was a small Southern fauna (species that do not occur <38ºS), comprising 15 % of the fauna, that reached a peak on the heavily sampled Tasmanian seamounts. These are largely subantarctic species, which in many cases are widespread throughout the Southern Ocean. Only a few of these species persisted onto the eastern slope of Tasmania. Finally there was a number of species (28 %) that were apparently endemic to the continental slopes of eastern and western Tasmania and Bass Strait. However, many of these species were either undescribed or rare, and the actual proportion of endemism is likely to be much less than this figure with further study. 2.6. Eastern Tasman Rise Unfortunately there were few data from the Eastern Tasman Rise. Most of the samples were deep (<2000 m). The fauna at this depth was broadly similar to that off the eastern slope of Tasmania. 2.7. Comparisons between taxonomic groups The different taxonomic groups exhibit a different pattern than the entire dataset. This is best examined by rank correlating the similarity matrices that underlie the ordinations and cluster diagrams (Clarke & Warwick 1994). No significance is attached to the correlations, the resulting coefficients are only used as an indicator that two patterns are similar. Coefficients of 1.0 indicate a perfect match between the entire dataset and a selected taxonomic group. Coefficients nearing 0 indicate a poor 14 correspondence between the patterns. Coefficients nearing -1 indicate a negative relationship, ie the patterns are reversed. The correlation coefficients are presented in Tables 3 and 4. Table 3 is based on an analysis of all 26 cells of 0.5 degree latitude that occur in the SE region (from 35ºS to 48ºS). Decapods show high correlations (R>0.5) for shallower depths, and ophiuroids are highly correlated with the overall pattern at deeper depths. Holothurians and pycnogonids in general did not correlate strongly with the entire dataset. This is due to the fewer specimens collected from these groups. Cells that do not contain specimens from all four taxonomic groups were excluded from the dataset and the analysis repeated (Table 4). With a few exceptions, the correlation coefficients were relatively high for all taxonomic groups at all depth strata that contained more than one cell of latitude with all groups present. This indicates that each group exhibits broadly similar geographic patterns when enough data is present. Slope East East East East East East West West West West West West Depth (m) No cells in analysis 200-500 26 500-1000 26 1000-1500 26 1500-2000 26 2000-2500 26 2500+ 26 200-500 26 500-1000 26 1000-1500 26 1500-2000 26 2000-2500 26 2500+ 26 Decapods 0.93 0.96 0.89 0.47 0.45 0.51 0.86 0.88 0.85 - Holo- Ophiur- Pycnothurians oids gonids 0.90 0.12 0.37 0.04 -0.42 0.51 0.71 0.06 -0.37 - 0.16 0.99 0.99 0.99 0.84 0.99 0.90 0.88 0.88 - -0.09 0.07 0.00 0.18 -0.42 0.51 0.86 -0.33 -0.36 - Table 4. Spearman Rank correlation coefficients between the entire dataset and selected taxonomic groups for each depth strata on either side of the SE region. The number of cells of latitude is kept constant even if not all taxonomic-groups were present. 15 Slope East East East East East East West West West West West West Depth (m) No cells in analysis 200-500 13 500-1000 14 1000-1500 10 1500-2000 9 2000-2500 1 2500+ 0 200-500 0 500-1000 5 1000-1500 0 1500-2000 0 2000-2500 0 2500+ 0 Decapods 0.93 0.92 0.98 0.94 0.72 - Holo- Ophiur- Pycnothurians oids gonids 0.80 0.64 0.90 0.79 0.42 - 0.95 0.92 0.96 0.94 0.52 - 0.50 0.84 0.90 0.84 0.72 - Table 5. Spearman Rank correlation coefficients between the entire dataset and selected taxonomic groups for each depth strata on either side of the SE region. Only the cells of latitude that contained all taxonomic groups are included in the analysis. 2.8. Change in distribution with depth Species richness of the echinoderms (ophiuroids and holothurians only), pycnogonids and decapods at 100 m depth intervals is shown in Figure 11a. This was the sum of samples collected in the SE region from 20 to 3500 m.. This graph showed that maximum species richness occurs on the shelf, followed by smaller peaks at 500 and 1000 m, then a rapid decline to the lower slope, with minor peaks 1400 and 2500 m. However much of this curve was directly related to collection effort. There were 331 samples from the upper shelf (<100 m), falling to 50 at 600 m.. Below 1900m there were less than 10 samples per 100 m (Figure 13). Different curves were obtained when samples collected by different methods were separated out (Figure 12). A lack of dredge/epibenthic sled samples from the upper slope was related to the decline of species richness from the shelf to 1000 m.. The minor peak at 400-600 m was related to the increase in trawl samples and the peak at 2500 to additional sled samples, which collected a large number of ophiuroids (Figure 11b). Not all taxonomic-groups had the same curves. The ophiuroids show an increase in diversity from the shelf to 1000 m (Figure 11 b), despite the decline in collection effort. The turnover in species was most pronounced from the shelf to 300 m (Figure 13). Whether this is an abrupt discontinuity or gradual change cannot be determined because of the low number of samples in the 150-300 m depth range. Other peaks in turnover (at 400 m, 1000 m and 2500 m) appeared to be related to species richness and collection efforts at these depths. The stacked plots of species richness also showed how species that can tolerate shallow depths rapidly decline with depth (Figure 11). Only 15% of shelf species were still present at 500 m, however one persisted to over 16 2600 m.. The upper to middle slope (to 2000 m) fauna frequently had large bathymetric ranges. A different fauna appears around 2500 m. The effect of depth on community structure is shown in figure 14 - a cluster diagram of epibenthic sled samples from the SE region. The two major clusters were based on shelf (0-210 m) and slope (> 200 m) samples. Shelf samples separated into an upper shelf and two lower-shelf clusters. The slope cluster separated into upper slope (<1000 m) and lower slope (> 1000 m) samples from the continental slope, and a separate cluster from the Tasmanian seamounts. The upper slope samples were further divided into a right cluster based on Southern Surveyor (2000) samples and a left cluster based on Franklin (1986) samples, a difference at least partially due to differences in collection gear. The Franklin material included many smaller species indicating the use of a smaller mesh size on the sled. In summary, after an initial rapid turnover of species from the shelf to slope, there was a gradual and continuous turnover of species until the lower slope (2500 m) after which there was a more pronounced faunal change. Communities were divided into shelf (<200 m), upper slope (200-1000 m), and lower slope (< 1000 m) categories. However, all these patterns were influenced by the current uneven collection effort. 2.9. Species endemic to the SE region Seventy-three continental slope species are apparently endemic to the SE region (Table 5). The majority of these species are undescribed (n=63) or described over the last 15 years (n=5). Many of these species are only known from the Tasmania seamounts. Not enough information is known about these animals to confidently assert their endemicity at this stage. Currently recognised deep-sea species are generally widespread, having ranges greater than the area of the SE region. However, recent genetic work indicates that some existing species are complexes of genetically distinct populations that may be considered distinct species with further research (O’Hara, unpublished). 17 Taxon Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Decapoda Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Family name Diogenidae Galatheidae Galatheidae Galatheidae Galatheidae Galatheidae Hippolytidae Hippolytidae Hippolytidae Hippolytidae Hippolytidae Hippolytidae Hippolytidae Paguridae Pasiphaeidae Polychelidae Sicyonidae Amphiuridae Amphiuridae Amphiuridae Amphiuridae Asteroschematidae Cucumariidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiacanthidae Ophiactidae Ophiomyxidae Ophiuridae Ophiuridae Ophiuridae Ophiuridae Ophiuridae Ophiuridae Ophiuridae Ophiuridae Genus Paguristes Munida Munida Munida Munida Munidopsis Eualus Lebbeus Lebbeus Leontocaris Leontocaris Leontocaris Thoralus Pagurid Pasiphaea Polycheles Sicyonia Amphioplus Amphioplus Amphiura Amphiura Asteroschema Species aciculus sp MoV 2672 sp MoV 2673 sp MoV 2674 sp MoV 2766 sp MoV 2677 sp MoV 2681 sp MoV 2679 sp MoV 2680 amplectipes bulga yarramundi sp MoV 2769 sp MoV 2683 kapala sp MoV 2719 australiensis sp MoV 2722 sp MoV 4529 sp MoV 4530 sp MoV 4531 sp. Amphicyclus Ophiacantha Ophiacantha Ophiacantha Ophiacantha Ophiacantha Ophiacantha Ophiocamax Ophiocamax Ophiocymbium Ophiomitrella Ophiomitrella Ophioplinthaca Ophiopristis Histampica Ophiobyrsa Ophiopyrgus Ophiura Ophiura Ophiura Ophiura Ophiura Ophiura Ophiurid mortenseni sollicita sp MoV 2731 sp MoV 4532 sp MoV 4536 sp MoV 4537 sp MoV 4541 applicatus sp MoV 4540 sp MoV 4539 sp MoV 2732 sp MoV 2779 sp MoV 2778 sp MoV 4535 sp MoV 4542 rudis sp MoV 4544 sp MoV 2728 sp MoV 3580 sp MoV 4523 sp MoV 4527 sp MoV 4528 sp MoV 4545 sp MoV 2733 Taxon Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Pycnogonida Family name Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Ammotheidae Austrodecidae Austrodecidae Callipallenidae Callipallenidae Callipallenidae Callipallenidae Callipallenidae Colossendeidae Nymphonidae Nymphonidae Phoxichilidiidae Phoxichilidiidae Phoxichilidiidae Phoxichilidiidae Phoxichilidiidae Phoxichilidiidae Rhynchothoracidae Genus Ammotheid Ascorhynchus Ascorhynchus Ascorhynchus Ascorhynchus Ascorhynchus Cilunculus Cilunculus Cilunculus Cilunculus Austrodecus Pantopipetta Callipallene Callipallene Callipallene Pallenopsis Pallenopsis Colossendeis Nymphon Nymphon Anoplodactylus Anoplodactylus Anoplodactylus Anoplodactylus Anoplodactylus Anoplodactylus Rhynchothorax Species sp slope 1 sp slope 2 sp slope 3 sp slope 4 sp slope 5 sp slope 6 sp slope 4 sp slope 6 sp slope 7 sp slope 8 sp slope 2 sp slope 1 sp slope 1 sp slope 2 sp slope 3 sp slope 1 sp slope 3 sp nov sp slope 3 sp slope 4 sp slope 2 sp slope 3 sp slope 5 sp slope 6 sp slope 7 sp. nov. sp slope 1 Table 5. Apparently endemic species from the continental slope of the SE region. 18 3. Macquarie Ridge 3.1. Methods The fauna of the Macquarie Ridge was analysed separately from the SE continental slope because of the differences in the taxa available for analysis and the lower overall collection effort. Data Most material was collected by BANZARE (1939), NZOI (1959-1965), Eltanin (1965-1968), ANARE (1986) and CSIRO Southern Surveyor (1999) cruises to the Macquarie Ridge. Material was collected using bottom trawls, except for a few epibenthic sleds collected by the 1999 Southern Surveyor Cruise. The dataset included echinoderms (all classes) and pycnogonids. In all there were 58 species in 246 collection (species/station) lots. Analysis The data were summarised into the presence/absence of each species in cells of one degree latitude along the Ridge (49-56ºS). The amount of longitude included varied with the width of the Macquarie Ridge at each latitude. Only material from the relatively narrow ridge was included in the analysis. Although the target depth segment was 200 m, not enough material was present at al longitudes to facilitate stratification of the dataset in this way. Nor was there enough material to analyse depths below 2000 m. Consequently, depth strata of 0200, 200-500, 500-1000, 1000-1500, 1500-2000 m were finally used in the final analysis. A species that occurs at the limit between depth strata (eg 500 m) was included in the next depth strata (500-1000 m). Two additional cells were created representing the northern (<=48ºS) and southern (>=57ºS) borders of the region based on whether each species is known to occur at more northern or southern latitudes. The data were sourced from published literature and collection databases. The actual data were supplemented by interpolated data, generated by recording a species as present for cells that are between known occurrences for each depth strata. The data were analysed using multivariate statistics, including MDS ordinations and cluster diagrams, using Bray-Curtis similarity coefficients and group-average linkage. The use of the Bray-Curtis similarity measure on presence/absence data is equivalent to using the Sorensen Binary coefficient. 19 Table 6. Geographic spread of the number of species recorded in each cell from the Macquarie Ridge (including interpolated records). Depth (m) 0-200 200-500 500-1000 1000-1500 1500-2000 2000-2500 2500+ 48+ 3 6 15 11 8 5 49ºS 50ºS 51ºS 52ºS 53ºS 54ºS 55ºS 56ºS 6 4 4 9 5 29 4 2 2 3 4 10 15 9 10 8 7 10 8 12 3 6 6 6 6 9 5 4 1 2 2 2 3 2 4 3 1 57+ 6 2 1 Table 7. Geographic spread of the number of collection stations by latitude (rows) and longitude (columns) from the Macquarie Ridge. Heavily surveyed cells include the Macquarie Island (54.5ºS) cell. Lat 49ºS 50ºS 51ºS 52ºS 53ºS 54ºS 55ºS 56ºS 150ºE 158ºE 159ºE 160ºE 162ºE 163ºE 164ºE 2 1 1 2 2 2 1 5 35 4 4 4 20 3.2. Results Depth strata 0-200 m Multivariate analysis Macquarie Island (54ºS) forms an outlier on the MDS. This is due to it being the only data rich cell at this depth in the analysis. Other cells are depauperate as there are few sections of the Macquarie Ridge with seafloor < 200 m. 200-500 m The data points are scattered across the MDS, reflecting the lack of systematic collections along the Macquarie Ridge at this depth (Table 6). 500-1000 m This graph (Figure 15a) shows a clear biogeographic gradient from the very north of the Macquarie Ridge, south of New Zealand (48ºS), to Macquarie Island (54ºS). The cells along the Southern Ridge are separate. Three groups can be distinguished using the cluster analysis (Figure 16): a) Northern Macquarie Ridge (48-52ºS) b) Central Macquarie Ridge, including Macquarie Island (5354ºS) c) Southern Macquarie Ridge (55-57ºS) 1000-1500 m The three groups identified for the 500-1000 m depth strata can also be clearly distinguished at this depth. 1500-2000 m The far north (48ºS) and far south cells (55-57ºS) can be distinguished from those along most of the Ridge. The animals present at this depth tend to be widespread Southern Ocean species. 3.3. Summary At depths where there is an adequate dataset along the entire Macquarie Ridge (5001500 m), the fauna separates into three geographic clusters, based on the northern (4852ºS), central (53-54ºS) and southern (55-57ºS) sections of the Ridge (Figures 15 and 16). This represents a biogeographic gradient from a New Zealand subantarctic fauna to an Antarctic fauna, with the central fauna representing a mixed zone with a few endemic species. At shallower depths (0-200, 500-1000 m), Macquarie Island becomes an outlier. This is partly due to the inconsistent distribution of shallow habitats along the Ridge and the presence of Macquarie Island endemic species. A similar widespread fauna occurs along most of the Ridge at the deepest depths analysed (1500-2000 m), although the far north (48ºS) and far south (55-57ºS) cells can be distinguished. 21 4. Conclusions This study must be considered a preliminary analysis of the benthic fauna of the SE Australian continental slope. The uneven sampling effort has been a major impediment to conclusive analysis. There are many more samples taken from the northern sections of the SE region, and around Macquarie Island along the Macquarie Ridge. There were very few samples from the South and Eastern Tasman Rises. This gradient of effort does affect the multivariate analyses. Nevertheless, there are some useful patterns in the data. The benthic fauna of the SE continental slopes is a gradient from a temperate fauna of the Tasman Sea to the subantarctic fauna of the South Tasman Rise. The temperate fauna persists as far as mid Tasmania (approximately 42ºS), and possibly will be found further south with increased collection effort. Subantarctic species become progressively more important in the region south of Tasmania. Many subantarctic species occur on the Tasmanian seamounts (44ºS) (Koslow et al. 2001). There is a large apparent difference between the eastern and western slopes, and much of this is due to the greater collection effort off the East Coast. Nevertheless, there is a small fauna (1.6%) that is only known from the western slope, and a larger (but as yet unquantified) fauna that is restricted to the Tasman Sea. There is a turnover of species with depth. There is a rapid turnover between the shelf fauna and the upper slope (150-300 m), followed by gradual change until 2500 m, after which there is a more pronounced faunal change. There are no samples from the abyssal plain. Communities were divided into shelf (<200 m), upper slope (200-1000 m) and mid-lower slope categories (>1000 m). The interaction between depth and latitude for the eastern slope was documented by Poore et al. (1994). The seamount fauna from Southern Tasmania is also distinct. There is a gradient along the Macquarie Ridge also, with the fauna of the Northern and Southern Ridges being distinct from each other and the fauna around Macquarie Island. The Northern Ridge supports subantarctic New Zealand species (O’Hara 1998) and the Southern Ridge Antarctic species. The shallow water fauna around the island is composed of shelf species from New Zealand and subantarctic species that have presumably arrived on kelp holdfasts from islands further west (O’Hara 1998). 22 4. Participants Dr Tim O’Hara (MV) identified most of the echinoderms; Dr Gary Poore (MV) and Dr. Shane Ahyong (AM) identified the decapods, David Staples (MV) the pycnogonids and Mark O’Loughlin (MV) the holothurians. Dr Tim O’Hara (MV) ran the analyses and compiled this document. 5. References Clarke, K.R. & Warwick, R.M. (1994). Change in marine communities: An approach to statistical analysis and interpretation. Natural Environment Research Council and Plymouth Marine Laboratory: UK. Koslow J.A., Gowlett-Holmes K., Lowry J.K., O´Hara T.D., Poore G.C.B., & Williams A. (2001). The seamount benthic macrofauna off southern Tasmania: community structure and impacts of trawling. Marine Ecology Progress Series 213: 111-125. O'Hara, T.D. (1998). Origin of Macquarie Island echinoderms. Polar Biology 20: 143151. O’Hara, T.D. and Poore, G.C.B. (2000). Distribution and origin of Southern Australian echinoderms and decapods. Journal of Biogeography 27: 1321-1335. Poore, G.C.B., Just, J. & Cohen, B.F. (1994). Composition and diversity of Crustacea Isopoda of the southeastern Australian continental slope. Deep-Sea Research 41: 677-693. 23 Eastern slope - 200-500 m 0 40 60 Eastern slope - 200-500 m Stress: 0.01 48+ 35+ 36-0 36-5 37-0 37-5 47-5 47-0 46-5 46-0 45-5 45-0 44-5 44-0 43-5 38-5 38-0 39-0 40-0 40-539-5 41-0 41-5 43-0 42-0 42-5 Figure 2. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 200-500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 24 43-5 43-0 47-5 47-0 46-5 46-0 45-5 45-0 44-5 48+ 44-0 38-0 37-5 39-0 38-5 39-5 41-5 41-0 40-5 40-0 42-5 42-0 36-5 100 36-0 80 35+ 37-0 Similarity 20 Eastern slope - 500-1000 m 20 60 Eastern slope - 500-1000 m Stress: 0.02 38-0 37-5 37-0 42-0 41-5 38-5 39-5 41-0 39-0 40-0 40-5 42-5 47-5 47-0 46-5 46-0 45-5 45-0 44-5 43-5 43-0 48+ 36-5 36-0 35+ 44-0 Figure 3. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 500-1000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 25 43-5 43-0 42-5 44-0 47-0 46-5 46-0 45-5 45-0 44-5 48+ 47-5 37-5 37-0 36-5 35+ 36-0 41-0 40-5 41-5 40-0 39-0 38-5 39-5 100 42-0 80 38-0 Similarity 40 Eastern slope - 1000-1500 m 20 60 Eastern slope - 1000-1500 m Stress: 0.01 44-0 42-5 43-542-0 43-0 41-5 38-5 41-0 40-5 40-0 39-5 39-0 46-5 46-0 45-5 45-0 44-5 38-0 47-0 47-5 48+ 37-0 36-0 37-5 36-5 35+ Figure 4. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 1500-2000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 26 37-0 36-0 37-5 35+ 36-5 39-5 39-0 41-0 40-5 40-0 41-5 38-5 42-0 43-5 43-0 42-5 38-0 48+ 44-0 47-5 47-0 45-0 44-5 46-5 100 46-0 80 45-5 Similarity 40 Eastern slope - 1500-2000 m 0 40 60 Eastern slope - 1500-2000 m Stress: 0.01 39-5 38-0 37-0 36-5 41-5 43-544-0 42-5 43-0 39-0 36-0 38-5 35+ 37-5 41-0 40-0 40-5 42-0 46-0 47-0 46-5 45-5 48+ 44-5 45-0 47-5 Figure 5. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 1500-2000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 27 40-5 40-0 41-0 39-0 39-5 41-5 38-5 43-0 42-5 43-5 42-0 37-0 36-5 37-5 35+ 36-0 38-0 48+ 44-0 47-5 47-0 46-5 46-0 45-5 100 45-0 80 44-5 Similarity 20 Eastern slope - 2000-2500 m 20 60 Eastern slope - 2000-2500 m Stress: 0.05 41-5 42-0 44-0 42-5 38-5 38-0 37-5 37-0 36-5 36-0 39-5 39-0 43-5 41-0 43-0 40-5 40-0 44-5 47-5 47-0 46-5 46-0 45-5 45-0 48+ 35+ Figure 6. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from 2000-2500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 28 39-0 38-0 37-5 37-0 36-5 36-0 39-5 38-5 40-5 40-0 41-0 43-5 43-0 35+ 42-5 42-0 41-5 44-0 47-0 46-5 46-0 45-5 45-0 100 44-5 80 48+ 47-5 Similarity 40 Eastern slope - 2500 m+ 20 60 Eastern slope - 2500 m+ Stress: 0.08 35+ 41-5 40-5 42-0 36-5 41-0 43-5 43-0 42-5 40-0 39-5 39-0 36-0 38-0 37-5 37-0 44-0 38-5 Figure 7. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the eastern SE Australian continental slope from below 2500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 29 43-0 42-5 43-5 41-0 39-5 39-0 40-0 44-0 40-5 38-5 42-0 41-5 38-0 37-5 37-0 36-0 100 36-5 80 35+ Similarity 40 Western slope - 200-500 m 0 40 60 Western slope - 200-500 m Stress: 0.01 41-5 48+ 41-0 40-0 40-5 39-5 35+ 36-5 36-0 39-0 38-0 37-5 37-0 38-5 43-0 42-0 42-5 Figure 8. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the western SE Australian continental slope from 200-500 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 30 42-5 42-0 43-0 41-5 36-5 36-0 37-0 38-0 37-5 35+ 38-5 40-5 39-5 40-0 41-0 100 39-0 80 48+ Similarity 20 Western slope - 500-1000 m 20 60 Western slope - 500-1000 m Stress: 0.05 35+ 36-0 36-5 47-5 47-0 46-5 46-0 45-5 45-0 44-5 48+ 37-5 37-0 38-0 43-0 42-5 42-0 43-5 41-5 39-5 39-0 40-0 40-5 41-0 38-5 44-0 Figure 9. Cluster analysis and MDS ordination of echinoderms, decapods and pycnogonids from the western SE Australian continental slope from 500-1000 m. Data consists of presence/absence of each species in cells of half a degree latitude and longitude. 31 43-0 42-5 42-0 43-5 47-0 46-5 46-0 45-5 45-0 44-5 48+ 47-5 44-0 39-5 39-0 40-0 40-5 38-0 37-5 37-0 41-5 41-0 38-5 100 36-5 80 35+ 36-0 Similarity 40 1 8 0 1 6 0 1 4 0 1 2 0 Numberofspecies 1 0 0 8 0 6 0 4 0 2 0 0 3 6 .0 3 7 .0 3 8 .0 3 9 .0 4 0 .0 4 1 .0 4 2 .0 4 3 .0 4 4 .0 4 5 .0 4 6 .0 4 7 .0 3 6 .5 3 7 .5 3 8 .5 3 9 .5 4 0 .5 4 1 .5 4 2 .5 4 3 .5 4 4 .5 4 5 .5 4 6 .5 4 7 .5 S ths p p N ths p p E n d e m ic W id e s p re a d a) Eastern slope 4 5 4 0 3 5 3 0 Numberofspecies 2 5 2 0 1 5 1 0 5 0 3 6 .0 3 7 .0 3 8 .0 3 9 .0 4 0 .0 4 1 .0 4 2 .0 4 3 .0 4 4 .0 4 5 .0 4 6 .0 4 7 .0 3 6 .5 3 7 .5 3 8 .5 3 9 .5 4 0 .5 4 1 .5 4 2 .5 4 3 .5 4 4 .5 4 5 .5 4 6 .5 4 7 .5 S ths p p N ths p p E n d e m ic W id e s p re a d L a titu d e b) Western slope Figure 10. Stacked plot of the number of species per grid. 32 1 4 0 1 2 0 1 0 0 noofspecies 8 0 6 0 m ind e p th(m ) o fe a c hs p e c ie s 4 0 > =2 5 0 0 <2 5 0 0 2 0 <1 5 0 0 <1 0 0 0 0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 2100 2200 2300 2400 2500 2600 2700 2800 2900 3000 3100 3200 3300 0 <5 0 0 <2 0 0 d e p th(m ) a) Decapods, pycnogonids, ophiuroids, holothurians 5 0 4 0 noofspecies 3 0 2 0 m ind e p th(m ) o fe a c hs p e c ie s 1 0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 2100 2200 2300 2400 2500 2600 2700 2800 2900 3000 3100 0 0 > =2 5 0 0 <2 5 0 0 <1 5 0 0 <1 0 0 0 <5 0 0 <2 0 0 D e p th(m ) b) Ophiuroids only Figure 11. Stacked plot of the species richness for each 100 m depth strata for all stations across the SE region. 33 1 4 0 1 2 0 1 0 0 noofspecies 8 0 6 0 4 0 2 0 100200 0 400500300 700800600 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 2100 2200 2300 2400 2500 2600 2700 2800 2900 3000 3100 3200 3300 3400 3500 0 A ll D re d g e T ra w l O th e r d e p th(m ) Figure 12. Species richness from various collection methods for 100 m depth strata across the SE region. 2 2 5 3 5 0 2 0 0 3 0 0 1 7 5 2 5 0 1 2 5 2 0 0 1 0 0 1 5 0 noofstations noofspecies 1 5 0 7 5 1 0 0 5 0 5 0 2 5 0 100200 0 300400 500600 700800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 2100 2200 2300 2400 2500 2600 2700 2800 2900 3000 3100 0 S p ptu rn o v e r(L ) S p pric h n e s s(L ) E ffo rt (R ) d e p th(m ) Figure 13. Species richness, species turnover and effort (no of samples) for 100 m depth strata across the SE region. 34 Figure 14. Cluster diagram of all samples from the SE region collected using an epibenthic sled (species abundance data is square root transformed). 0 20 40 60 80 10 0 < 130 m (Bass Strait) 1000-3000 620-1900 m (slope) m (seamounts) 150-1000 m (slope) 130-210 m (Outer shelf) (mixed outliers) 35 0-200 m 1000-1500 m Stress: 0.03 Stress: 0.01 55 49 56 48+ 5051 53 52 5049 51 55 54 57 + 53 48+ 52 54 200-500 m Stress: 0.01 1500-2000 m Stress: 0 48 + 48+ 53 54 52 51 50 49 55 57+56 54 52 50 49 51 53 500-1000 m Stress: 0 48+ 5657 + 49 50 51 55 52 Figure 15. MDS ordinations of species from the Macquarie Ridge, based on the presence/absence of each species along the Ridge in cells of one-degree latitude. Macquarie Island is located at 54ºS. 53 54 36 500-1000 m 0 20 40 Bray-Curtis Similarity 60 80 100 55 56 57+ 53 54 48+ 52 49 50 51 Figure 16. Cluster diagram of species from the Macquarie Ridge, based on the presence/absence of each species along the Ridge in cells of one-degree latitude between 500-1000 m. Macquarie Island is located at 54ºS 37