A comparison of megafaunal biodiversity in two contrasting submarine canyons on Australia’s

southern continental margin

David R. Currie and Shirley J. Sorokin

SARDI Publication No. F2010/000981-1 SARDI Research Report Series No. 519

SARDI Aquatic Sciences PO Box 120 Henley Beach SA 5022

February 2011

Report to the South Australian Department of Environment and Natural Resources

A comparison of megafaunal biodiversity in two contrasting submarine canyons on Australia’s southern continental margin

Report to the South Australian Department of Environment and Natural Resources

David R. Currie and Shirley J. Sorokin

SARDI Publication No. F2010/000981-1 SARDI Research Report Series No. 519

February 2011 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

This Publication may be cited as: Currie, D.R and Sorokin, S.J (2011). A comparison of megafaunal biodiversity in two contrasting submarine canyons on Australia’s southern continental margin. Report to the South Australian Department of Environment and Natural Resources. South Australian Research and Development Institute (Aquatic Sciences), Adelaide. SARDI Publication No. F2010/000981-1. SARDI Research Report Series No. 519. 49pp.

South Australian Research and Development Institute SARDI Aquatic Sciences 2 Hamra Avenue West Beach SA 5024

Telephone: (08) 8207 5400 Facsimile: (08) 8207 5406 http://www.sardi.sa.gov.au

DISCLAIMER The authors warrant that they have taken all reasonable care in producing this report. The report has been through the SARDI Aquatic Sciences internal review process, and has been formally approved for release by the Chief, Aquatic Sciences. Although all reasonable efforts have been made to ensure quality, SARDI Aquatic Sciences does not warrant that the information in this report is free from errors or omissions. SARDI Aquatic Sciences does not accept any liability for the contents of this report or for any consequences arising from its use or any reliance placed upon it.

© 2011 SARDI This work is copyright. Apart from any use as permitted under the Copyright Act 1968 (Cth), no part may be reproduced by any process, electronic or otherwise, without the specific written permission of the copyright owner. Neither may information be stored electronically in any form whatsoever without such permission.

Printed in Adelaide: February 2011

SARDI Publication No. F2010/000981-1 SARDI Research Report Series No. 519

Author(s): Dr David R. Currie and Ms Shirley J. Sorokin

Reviewer(s): Dr Nathan J. Bott and Ms. Sonja L. Hoare

Approved by: Dr Jason E. Tanner Principal Scientist – Marine Environment & Ecology

Signed:

Date: 1 February 2011

Distribution: South Australian Department of Environment and Natural Resources, SAASC Library and University of Adelaide Library

Circulation: Public Domain

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TABLE OF CONTENTS

LIST OF FIGURES ...... III

LIST OF TABLES...... III

LIST OF APPENDICES ...... IV

EXECUTIVE SUMMARY ...... 1

1 INTRODUCTION ...... 3

1.1 BACKGROUND...... 3 1.2 OBJECTIVES ...... 3

2 METHODS...... 5

2.1 FIELD SAMPLING ...... 5 2.2 LABORATORY PROCESSING ...... 6 2.3 STATISTICAL ANALYSES...... 6 2.3.1 Univariate patterns...... 6 2.3.2 Environmental parameters ...... 6 2.3.3 Multivariate patterns ...... 6

3 RESULTS...... 8

3.1 ENVIRONMENTAL CHARACTERISTICS ...... 8 3.1.1 Bathymetry...... 8 3.1.2 Sediment composition and structure...... 8 3.1.3 Oceanography...... 9 3.2 EPIFAUNAL COMPOSITION ...... 9 3.3 SPATIAL PATTERNS IN SPECIES RICHNESS AND BIOMASS ...... 10 3.4 ENVIRONMENTAL LINKAGES - RICHNESS AND BIOMASS ...... 10 3.5 EPIFAUNAL COMMUNITY STRUCTURE ...... 11 3.6 ENVIRONMENTAL LINKAGES – COMMUNITY STRUCTURE ...... 12

4 DISCUSSION ...... 13

REFERENCES ...... 17

AKNOWLEDGEMENTS ...... 21

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LIST OF FIGURES Figure 1. Map showing the locations of Bonney and du Couedic canyons (unfilled rectangles) on the south-east continental margin. Contour lines presented follow 100 m depth intervals. .. 22

Figure 2. Bathymetric map of du Couedic Canyon showing the locations of 5 survey sites sampled for macro-epibenthos, sediments and near-bed water properties. Labels denote location relative to the main canyon axis (DCC = du Couedic Centre) and sampling depth in metres (100, 200, 500, 1000, 1500)...... 23

Figure 3. Bathymetric map of Bonney Canyon showing the locations of 6 survey sites sampled for macro-epibenthos, sediments and near-bed water properties. Labels denote location relative to the main canyon axis (BC = Bonney Centre) and sampling depth in metres (100, 200, 500, 1000, 1500, 2000)...... 24

Figure 4. Plots of (a) mean wet weight, (b) total number of species of each major phylum collected during the survey, and (c) total number of sites (out of 11) at which specimens belonging to each major phylum were collected. Values for each variable are shown as percentages above each bar...... 25

Figure 5. Total (a) number of species, and (b) wet weights of macro-epibenthos collected during sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons. 26

Figure 6. Dendrogram and non-metric MDS ordination of epifaunal community structure at 11 depth-stratified sampling stations at Bonney and du Couedic canyons. Sampling depths are represented by different symbols (upright triangle = 100 m, diamond = 200 m, circle = 500 m, inverted triangle = 1000 m, square = 1500 m, plus = 2000 m) and canyon locations by different shades of fill (black = Bonney, white = du Couedic). Three station groupings are identified at the 5% Bray-Curtis similarity level (dotted line): shelf (100-200 m), upper slope (500 m) and mid- slope (1000-2000 m)...... 27

LIST OF TABLES Table 1. Summary of environmental factors at 11 epibenthic sampling sites at Bonney and du Couedic canyons...... 28

Table 2. Spearman’s rank correlation coefficients between epifaunal richness and biomass and adjacent environmental conditions at 11 depth-stratified sampling stations at Bonney (B) and du Couedic (dC) Cannon. Significant positive (+) and negative (-) correlations are denoted at the ** 1% and * 5% level...... 29

Table 3. Mean biomass (kg per ha ± s.e.) of macro-epibenthic species in three station groups identified from MDS classification. Species listed were identified from SIMPER analyses as contributing ≥5% to the similarity within and dissimilarity between regional groupings. Those species indicative of each regional grouping (contributing ≥10% to the total similarity within a group) are highlighted in bold. Species are ranked in order of decreasing biomass across all station groupings...... 30

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LIST OF APPENDICES Appendix 1. Taxonomic and functional classification of 184 macro-epibenthic species collected from sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons. All species codes given here refer to material lodged in the South Australian Museum...... 31

Appendix 2. Photographic plates depicting 184 organisms collected in epibenthic sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons...... 35

Appendix 3. Summary list of species biomasses (kg per hectare) collected during sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons...... 47

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EXECUTIVE SUMMARY

1. This report describes the composition, distribution and biomass of macro-epibenthic organisms along the central axes of two South Australian submarine canyons with contrasting topographies (i.e. the v-shaped shelf-breaking du Couedic Canyon, and the box shaped upper-slope Bonney Canyon).

2. Quantitative samples of epibenthos were collected from 5 depth-stratified sampling sites at du Couedic Canyon (100, 200, 500, 1000 & 1500 m), and 6 depth-stratified sampling sites at Bonney Canyon (100, 200, 500, 1000, 1500 & 2000 m), using a 1 m wide sled which was towed across the seabed for 500 m.

3. A total of 184 putative species representing 11 phyla were collected. Less than 10% of these taxa (16/184) could be confidently assigned to existing species, and it appears that a large proportion of the canyon fauna is undescribed.

4. Sessile suspension-feeding organisms (primarily poriferans, molluscs, cnidarians and bryozoans) dominated samples, and collectively comprised over 97% of the biomass and 77% of the species richness. The most common free-living organisms were echinoderms which comprised 2% of the biomass and 13% of the species collected.

5. Species numbers broadly declined with increasing depth in both canyons, however species richness was more than twice as high at du Couedic Canyon (140 sp) than it was at Bonney Canyon (60 sp).

6. Epifaunal biomass was also concentrated inshore in the shallowest depths of both canyons, but was over an order of magnitude higher at du Couedic Canyon than at Bonney Canyon (e.g. 1057 vs. 22 kg per ha at 100 m depth).

7. Bivariate correlation analyses highlighted a number of strong abiotic relationships (e.g. between depth, temperature and mud content), but linkages to species richness and biomass were not consistent between the two canyons.

8. Cluster analyses of species biomass data revealed a strong environmental gradient running perpendicular to the coast in both canyons, and highlighted a progressive shift in species composition.

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9. Three station groupings characterised by small groups of species with narrow distributions were identified on the shelf, upper slope and mid slope. These community groupings were largely explained by depth and dissolved oxygen concentration.

10. Spatial patterns in epifaunal community structure were broadly consistent with depth-related discontinuities in water circulation along Australia’s southern continental margin.

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1 INTRODUCTION

1.1 Background

Submarine canyons are becoming increasingly recognised as sites of enhanced productivity in the oceans (Genin, 2004), often vitally important to sustaining fish production. Studies in the Mediterranean, North Atlantic and Pacific Ocean have shown that canyons generate complex flows, the net result of which can be higher regional productivity (Allen et al., 2001; Bosley et al., 2004; Greene et al., 1988; Macquart-Moulin and Patriti, 1996). Off the Bonney Coast and Kangaroo Island, numerous canyons exist, that may provide nutrient and sediment paths between the deep ocean slope and coast (Hill et al., 2005). During summer, westward currents over the slope generated by the Flinders Current and meso-scale eddies are thought to produce currents within the canyons sufficient to move sediments and nutrients up slope (Griffin et al., 1997; Lewis, 1981; McClatchie et al., 2006; Middleton and Platov, 2003; Nieblas et al., 2009; Schahinger, 1987). This may result in hotspots of pelagic and benthic productivity and biodiversity. The Bonney Coast and Kangaroo Island canyons are of particular importance because the area is being explored for hydrocarbons (Boult et al., 2006). Tar balls stranded along the South Australian coast may be transported up the canyons from natural leaks at the base of the slope. If proven, this may indicate oil-bearing sediments buried in up to 4000m deep water, in this un-drilled region. Before a need arises to manage hydrocarbon extraction, it is imperative to understand the importance of these canyons to regional productivity.

During February 2008, the National Facility RV Southern Surveyor undertook a dedicated research voyage to assess the oceanographic and biological significance of two canyon systems off South Australia (Bonney and du Couedic; Figure 1) (Currie, 2008). One of the components of this multidisciplinary study involved the collection in sled tows of almost 150 kg of biota from the seafloor at 11 depth-stratified (100 – 2,000 m) sites located along the central axes of these two canyons. These faunal collections were frozen immediately after collection and offer an unprecedented insight into the composition, biomass and distribution of mega- benthos within these remote and poorly studied habitats.

1.2 Objectives

This report presents the results of the first quantitative survey of the epibenthic assemblages of Bonney and du Couedic canyons, undertaken during 2008. The objectives of the study were: (1) to provide a synoptic picture of the abundance, biomass and diversity of the macro- epibenthic fauna at the two canyon systems; (2) to assess the relative contributions of ambient

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environmental conditions on observed distributional patterns; and (3) to evaluate patterns in community structure in relation to canyon physiography and regional biodiversity.

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2 METHODS

2.1 Field sampling

Initially, a broad-scale mapping survey of each canyon was conducted using multi-beam sonar (Kongsberg-Simrad EM300). This provided high-resolution bathymetry of the target sites, and subsequently guided our detailed sampling of the area.

Quantitative samples of epifauna were collected using a 1 m wide Ockleman sled fitted with a 5 cm mesh bag. This sled was towed over a 500 m distance at each of 5 depth-stratified sampling stations at du Couedic Canyon (DC_100, DC_200, DC_500, DCC_1000, DC_1500; Figure 2) and 6 depth stratified sampling stations at Bonney Canyon (BC_100, BC_200, BC_500, BC_1000, BC_1500, BC_2000; Figure 3). On retrieval the catch was emptied on deck and a photograph taken. The collective weight of each shot was then determined and the catch transferred to labelled plastic bags before being snap-frozen. A maximum sub-sample of 35 kg of the homogenised catch was retained for laboratory analysis from each sled shot, while the remaining catch was discarded overboard.

The composition and structure of the seabed at each sampling site was determined from replicate 0.1 m² Smith-McIntyre grab samples. Two sediment sub-samples (70 ml and 10 ml) were collected from each grab by scraping an open vial across the top of each sample. These were snap-frozen and stored at -20 ºC before being analysed. The larger of the two sediment sub-samples was wet-sieved through an agitated stack of Endicott sieves to determine the grain-size structure and sorting coefficients of the sediments. The smaller sample was freeze dried, sieved, and then ground to a talcum-powder consistency before being processed in an elemental analyser (LECO TruSpec CN) to determine organic carbon and total nitrogen content.

Profiles of water temperature, salinity and pressure were recorded at all sampling sites (except BC_2000) using a Seabird SBE911 CTD (Conductivity, Temperature, Depth) unit fitted with modular sensors for dissolved oxygen (Aanderaa Optode 3975) and fluorescence (Chelsea AQUAtracka). These instruments were attached to the vessel’s 24-bottle rosette frame, and lowered to within 20 m of the seabed during each cast. Seabird-supplied calibration factors were used to calculate pressure and temperature, and their accuracy validated over the course of the voyage by examining, and correcting, deviations in the sensor records before and after each cast (Beattie, 2008). A series of 18 Niskin bottles mounted on the rosette frame were used to collect water samples at up to 6 depths on each cast. These samples were primarily collected to estimate spatial variability in phytoplankton productivity, but were also used to calibrate the salinity, oxygen and fluorescence sensors. As this study focused on epibenthos,

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CTD data extracted from the deepest part of each vertical cast were utilised in all subsequent analyses.

2.2 Laboratory processing

Sled samples were defrosted and sorted in the laboratory to the lowest taxonomic level (species level where possible) before being counted and weighed. During this process, all broken shells and rocks were discarded, while fragments of the same non-unitary organism (e.g. colonial ascidians) were consolidated and collectively weighed and counted as a single entity. Voucher specimens of all species collected were photographed before being preserved in 70% ethanol. All voucher specimens have been lodged at the South Australian Museum, Adelaide.

2.3 Statistical analyses

2.3.1 Univariate patterns

One-way analysis of variance (ANOVA) was used to test differences in total species richness and biomass between the two canyon systems and among the different depth strata. Prior to conducting all ANOVAs, homogeneity of variance was examined using Levene’s test and heterogeneity removed using a log10(n+1) transformation.

2.3.2 Environmental parameters

Spatial trends in epibenthic biomass and species richness were examined in relation to the physical, chemical and biological attributes at each sampling station. The relative strength of each environmental relationship was assessed independently for each canyon using Spearman’s rank correlation coefficients.

2.3.3 Multivariate patterns

Depth and canyon-related differences in epibenthic community structure were examined using Bray-Curtis (B-C) dissimilarity measures (Bray and Curtis, 1957). A single square-root transformation was applied to the data before calculating the B-C dissimilarity measures. This transformation was necessary to prevent a small number of large species unduly influencing the B-C dissimilarity measures (Clarke, 1993).

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The computer package PRIMER (Primer-E Ltd., Plymouth) was used to generate B-C dissimilarities and to undertake all multivariate analyses (Clarke and Gorley, 2001). Spatial patterns in dissimilarity were initially mapped using a combination of hierarchical agglomerative clustering and non-metric multi-dimensional scaling (nMDS), and depth and canyon-related differences tested using a permutational analysis of variance (PERMANOVA) (Anderson, 2001). A similarity percentages (SIMPER) routine (Clarke and Gorley, 2001) was subsequently used to identify those species contributing most to observed differences. Finally, the extent to which measured environmental variables accounted for any community groupings was tested using a biological environmental (BIOENV) routine (Clarke and Ainsworth, 1993).

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3 RESULTS

3.1 Environmental characteristics

3.1.1 Bathymetry

Swath bathymetric data collected during the survey highlight marked differences in the seafloor topographies of the two canyons (Figures 1 & 2). At du Couedic (Figure 2), a prominent v- shaped gouge in the outer shelf (20 km in length) characterises the upper section of the canyon. Beyond this (>200 m depth), the seafloor is highly folded and deeply incised, with numerous narrow channels and chutes extending across the slope. Here, below 1500 m, the canyon floor drops abruptly and forms a spectacular corrie with near-vertical headwalls (>1 km in height). The main branch of the canyon continues below this feature as a narrow (2 km wide) gorge. Bonney Canyon, by comparison, does not extend onto the continental shelf, but has a well-defined headwall on the upper slope (~800 m depth; Figure 3). Below the headwall, the canyon is narrow (3 km wide) and deeply incised (>1 km deep) with steep sidewalls (gradient > 1:1). The floor of the canyon is terraced and bears numerous scallop-shaped scars, indicative of erosion and slumping.

3.1.2 Sediment composition and structure

Samples of sediment taken within the two canyons were variable in structure and ranged from silt, to coarse-sand and gravel (Table 1). These sediments were found to be composed almost entirely of biogenic material, including fragments of sponges, bryozoans, molluscs, coralline algae and foraminifera. Spatial patterns in grain-size were broadly consistent with patterns in bathymetry. Sediments in both canyons were typically coarsest in the shallow inshore waters, but became progressively finer with increasing depth and distance offshore. Sediment sorting, by comparison, was less clearly related to depth, and was greatest at the 1000 m station at Bonney Canyon (Table 1).

The organic carbon content of the sediments ranged between (<0.01 and 0.23%) at du Couedic Canyon, and (0.06 and 1.07%) at Bonney Canyon, and broadly reflected trends in sediment size structure (Table 1). Notably, organic carbon content was found to be lower in the coarser sediments of the shelf than in the muddier sediments of the slope. In particular, organic carbon was concentrated in those sediments occurring at depths of 1000 and 1500 m inside the central axis of Bonney Canyon. A similar distribution pattern was also observed for concentrations of sedimentary nitrogen (Table 1).

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3.1.3 Oceanography

Marked depth-related differences in near-bottom water temperatures were observed along the central axis of both du Couedic and Bonney canyons (Table 1). Near-bed water temperatures gradually declined with increasing depth at both canyons (11.21-2.87 ºC at du Couedic, 10.69- 2.66 ºC at Bonney), but were marginally lower at all comparative depth strata in the more southerly Bonney Canyon.

Salinity varied little at both canyons, and ranged between 34.39 and 35.00 at du Couedic, and between 34.39 and 34.91 at Bonney Canyon (Table 1). Patterns in salinity were not well matched with temperature, but were generally higher in the shallower inshore waters of both canyons, than in the deeper offshore waters. Dissolved oxygen concentrations, by comparison, were much more tightly matched to temperature (Table 1), and were higher (>240 µM/l) in the warmer, shallower, waters (>8 ºC, <500 m depth) of both canyons, and lower (<200 µM/l) in the cooler deeper waters (<6 ºC, >1000 m depth). Near-bottom chlorophyll concentrations were also elevated in the shallow inshore waters of both canyons, and ranged between 10.34 and 15.54 µg/l at du Couedic, and between 10.05 and 13.87 µg/l at Bonney Canyon (Table 1)

3.2 Epifaunal composition

A total of 184 species representing 11 phyla were collected from the 11 depth-stratified sampling sites (Appendix 1-3). Porifera (sponges) were the dominant phyla in terms of biomass, and accounted for 84% of the standardised catch (mean biomass χ B = 158.28 kg per ha) (Figure 4a). Molluscs (shellfish) and cnidarians (corals) were also relatively well

represented, and comprised approximately 7% ( χ B = 13.80 kg per ha) and 4% ( χ B = 7.66 kg per ha) of the standardised catch, respectively. All other phyla collected, including Annelida (bristle worms), Arthropoda (crabs), Brachiopoda (lamp shells), Bryozoa (lace corals), Chordata (fish), Echinodermata (starfish), Nemertea (ribbon worms) and Urochordata (sea squirts) were

much less common, and individually comprised less than 2% of the catch weight ( χ B < 3.74 kg per ha).

Porifera were also the best represented phylum in terms of species richness, and accounted for 47% (87) of the species collected (Figure 4b). Echinoderms, cnidarians and bryozoans were also well represented, and accounted for 13% (24), 12% (22) and 8% (15) of the total species richness. Most other phyla collected, including annelids, arthropods, brachiopods, chordates,

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molluscs, nemerteans and urochordates were relatively less diverse and represented by fewer than 5% (9) of the total species collected.

Echinoderms were the most widely distributed phyla and occurred at all 11 survey sites. arthropods, bryozoans, cnidarians, molluscs, poriferans and urochordates were also widespread, and found at between 72% (8) and 82% (9) of sites (Figure 4c). All other phyla collected, including annelids, chordates, brachiopods and nemerteans had much more restricted distributions, and were encountered at less than 36% (4) of all sites sampled.

3.3 Spatial patterns in species richness and biomass

Most of the faunal richness in both canyons was concentrated inshore at the shallowest depths (i.e. 100 m and 200 m), and broadly declined with increasing depth (Figure 5a). Species richness was higher at du Couedic Canyon than at Bonney Canyon, with 140 species collected from 5 sled shots at du Couedic, and just 60 epibenthic species collected from 6 sled shots at Bonney Canyon. Of these species, 123 were unique to du Couedic, 43 were only found at Bonney, while a further 17 species were collected at both canyons. Du Couedic Canyon also supported relatively higher numbers of species than Bonney Canyon at all of the shallower depth strata surveyed (i.e. 100 m, 200 m and 500 m; Figure 5). However, this difference did not hold at the deeper sites (i.e. 1000 m and 1500m), where richness was marginally higher at Bonney than at du Couedic. As a result of this interaction, ANOVA tests for differences in species richness between the two canyons were not significant (F (1, 9) = 1.70, p = 0.225).

Epifaunal biomass, like species richness, was also concentrated inshore at the shallowest depths in both canyons (i.e. 100 m and 200 m), but was more than an order of magnitude higher at du Couedic Canyon than at Bonney Canyon (100 m = 1057 vs. 22 kg per ha; 200 m = 941 vs. 28 kg per ha) (Figure 5b). Depth-related difference in biomass also mirrored those of species richness in the deeper sections of both canyons, with higher biomasses of epifauna being collected at depths of 1000 m and 1500 m inside Bonney Canyon. On account of this interaction, ANOVA tests for differences in biomass between the two canyons were not

statistically significant (F (1, 9) = 0.816, p = 0.390).

3.4 Environmental linkages - richness and biomass

No significant relationships were detected at Bonney Canyon between either biological parameter considered in this study (i.e. species richness and biomass) and any of the

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measured environmental factors (Table 2). At du Couedic, by comparison, species richness was positively correlated with both temperature and salinity, while the faunal biomass was positively correlated with temperature and negatively correlated with depth, % mud and the sedimentary carbon content (Table 2). There were also significant correlations at Bonney Canyon between the oxygen saturation of the near bed water and the sampling depth, temperature, salinity, % mud and sedimentary carbon content. Besides this, no other environmental parameter varied in relation to species richness or biomass.

3.5 Epifaunal community structure

Three discrete station groupings were separated at the 5% B-C similarity level (Figure 6). These included a “shelf” group comprising all 4 stations surveyed at depths of 100 m and 200 m inside Bonney and du Couedic canyons, an “upper slope” group containing the 2 stations surveyed at a depth of 500 m inside Bonney and du Couedic canyons, and a “mid slope” group consisting of all 5 stations surveyed at depths between 1000 m and 2000 m inside Bonney and du Couedic canyons.

SIMPER analysis was undertaken to determine which taxa contributed most to similarities within and differences between the three station groupings. Biomasses of the 16 species contributing ≥5% to within-group similarity or between-group dissimilarity for at least one of the three groupings are given in Table 3. Results from the SIMPER analysis indicate that all station groupings are characterised by small subsets of species with restricted distributions.

The “shelf” group was the most diverse and consisted of 123 species, 114 (93%) of which were found only at stations located in depths of 200 m or less. Three species of Porifera and one species of Cnidaria typified this group, and contributed more than 10% to the within-group similarity. These included the slimy orange sponge Spheciospongia sp., the brown fan-shaped sponge Chalinid sp., the massive yellow sponge Polymastia sp. 2, and the branching brown hydroid Clathrozoon wilsoni.

The “upper slope” group was the least diverse and was composed of 24 species. Of these, 17 (71%) were unique to stations sampled at 500 m depth. This group was characterised by just one species, the burrowing heart urchin Brissus sp. This small deposit-feeding echinoderm collectively accounted for 12% of the total group biomass, and 98% of the within-group similarity.

The “mid slope” group contained 46 species, 43 (93%) of which were found exclusively at stations located in depths between 1000 and 2000 m. Two echinoderm species, the starfish

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Astropecten sp. 1 and the sea cucumber Holothuroid sp. 2, dominated the catches from this depth and accordingly characterised this station grouping.

3.6 Environmental linkages – community structure

The PRIMER routine BIOENV was used to assess the correspondence and significance of environmental data from the seafloor to the three station groupings identified from the community analyses. Measures of temperature and % mud were excluded from these analyses as they are highly correlated and co-varied with depth. The best fit was with depth (ρw = 0.64),

which in combination with the dissolved oxygen concentration gave a best fit of ρw = 0.71. The remaining variables (salinity, chlorophyll concentration, sediment sorting, sediment carbon and sediment nitrogen) were individually much more weakly correlated with the community structure

(ρw < 0.40), and failed to provide an improved explanation for the biological pattern when included with depth in a stepwise procedure.

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4 DISCUSSION Most benthic communities at depths below the photic zone are dependent on sinking water column production as a major source of food, hence the quality and quantity of organic matter reaching the seafloor is an important influence on benthic community structure and biomass (Smetacek, 1984). While hotspots of benthic diversity may be highly correlated with regions of enhanced surface phytoplankton concentrations, standing-stock and surface production data are not always concurrent (Johnson et al., 2007). Horizontal advection can complicate this linkage through the transport of sinking phytoplankton to a bottom area that is distant from the surface waters where they were abundant (Lampitt et al., 1995). Decoupling between herbivory and primary production can further modify the export of pelagic production to the benthos as a result of changes in zooplankton grazing rates (Ambrose and Renaud, 1995). The extent to which such factors influence epibenthic distributions around Bonney and du Couedic canyons is uncertain, however it is notable that faunal biomasses and richness were generally higher on the shallow shelf waters where benthic chlorophyll levels were elevated.

Chlorophyll concentrations on shelf waters may be enhanced by upwelling of cold nutrient-rich waters that support increased phytoplankton productivity. Off South Australia, seasonal upwelling is regionally significant along the continental margin, and is understood to promote discrete hotspots of pelagic productivity off the lower Eyre Peninsula, Kangaroo Island, and the Bonney coast (Ward et al., 2006). These hotspots of productivity may well be linked to variations in shelf-break topography, which could also account for the large differences in faunal diversity and biomass between Bonney and du Couedic Canyon. Alternatively, the large discrepancy in biomass and richness between du Couedic and Bonney Canyon may stem from the former canyon’s unique location relative to the Spencer Gulf. During summer, evaporative forcing causes the head waters of Spencer Gulf to become hypersaline (Lennon et al., 1987). As these waters cool during the Austral winter, the high-salinity water becomes dense enough to form an outflowing bottom current. This current, known as Bonaparte’s Tongue, flows out of the gulf and across the Lincoln Shelf towards du Couedic Canyon, bringing with it high quantities of particulate organic matter (Smith and Veeh, 1987). As sponges and other filter feeding organisms (e.g. ascidians, bryozoans) are dependant on particulate organic matter as a source of food (Diaz and Klaus, 2001), it is suggested that this mechanisms may be responsible for the exceptionally high biomasses and diversities of sponges and other suspension feeding epibenthos at the head of du Couedic Canyon.

While faunal richness and biomass broadly declined in both canyons with increasing depth, marked shifts in community structure were also observed in relation to depth. Such bathymetric changes in community structure are widely reported on shelf and slope habitats (Cartes et al.,

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2009; Kroncke et al., 2003; Louzao et al., 2010), but geographical differences between studies, as well as variations in the range of depths considered or the classification methods employed, mean that patterns are often contradictory. In our study spanning the shelf and slope (100-2000 m) off southern Australia, three distinct communities were identified. These included a diverse “shelf” community (100-200 m) characterised by several species endemic to southern Australia (e.g. the cnidarian Clathrozoon wilsoni, the poriferan Spheciospongia papillosa, and mollusc Pyxipoma weldii), a low diversity “upper slope” community (500 m) typified by more widely- ranging southern temperate species (e.g. the cnidarians Caryophyllia planilamellata and Desmophyllum dianthus), and a moderately diverse “mid slope” community (1000-2000 m) that includes species with circumglobal distributions (e.g. the mollusc Lucinoma galatheae). These results appear to be broadly consistent with demersal fish data from a narrower depth range off southern Australia (500-1200 m) (Koslow et al., 1994); where distinct assemblages are apparent in the upper (500 m) and mid slope (800-1200 m).

Koslow et al. (1994), in recognising affinities between the southeast Australian mid slope fish communities and those at similar depths in the North Atlantic, suggested that biogeographic patterns were consistent with ocean circulation at intermediate depths. Notably, they observed that their mid slope community resided within the core depth range (800-1200 m) of the Antarctic Intermediate Water mass, which extends around the northern rim of the Southern Ocean (Fine, 1993). Like Koslow et al. (1994), our mid slope community also corresponds with Antarctic Intermediate Water, but faunal discontinuities at the upper slope and shelf also coincide with estimates of the upper and lower vertical boundaries (400-900 m) of the westward moving Flinders Current (Middleton and Bye, 2007). Similar zonational patterns in demersal fish are also reported on the West Australian continental slope, where community breaks at 300 m and 700 m depth coincide with lower boundary limits of the near-surface Leeuwin Current and upper extent of the Antarctic Intermediate Water, respectively (Williams et al., 2001). While such correlations are not necessarily causative, they are intriguing, and suggest that ocean circulation patterns play an important role in structuring benthic communities at regional scales (10-1000 km).

In our community analyses, depth was identified as the most influential factor structuring epibenthic assemblages in Bonney and du Couedic Canyon. However, depth is unlikely to be the primary causal factor determining faunal composition. This is because many other physical/chemical variables co-vary with depth (e.g. temperature, salinity, oxygen) and may also influence the distribution of benthic species. Water circulation patterns for example, which may also vary with depth, can influence benthic communities in several ways. In particular, water circulation can modify other water column processes, such as near-bottom flow, that bring food and new recruits to the community (Snelgrove and Butman, 1994). They can also influence the

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physical heterogeneity of the seafloor through processes of erosion and deposition, and thereby directly influence the distribution of habitats and associated species (Hall, 1994). A range of other biological processes (e.g. predation, competition) are also likely to have important influences on the distribution and abundance of macrobenthic species, particularly at local scales (1-1000 m), but remain unmeasured.

Although the 184 putative species collected from Bonney and du Couedic Canyon represent only a small component of Australia’s marine macrofaunal biodiversity (>250,000 species (Butler et al., 2010)), the species richness is relatively high when compared with other shelf and slope environments. For example, a total of 196 epifaunal species were collected from more than ten times as many sled tows (152) in the North Sea (70-200 m)(Basford et al., 1990). Relatively few species (140) were also collected from more than five times as many sled tows (65) in the north-western Atlantic (50-110 m) (Thouzeau et al., 1991). A total of 181 epifaunal species were collect from a marginally larger number of tows (18) during a study in the eastern Atlantic (425-1050 m) (Sanchez et al., 2008). While in a survey off seamounts in the Southern Ocean, 242 putative species were collected from 34 trawl shots (660-1700 m) (Koslow et al., 2001). On the basis of these preliminary comparisons it is tempting to suggest that epifauna diversity from Bonney and du Couedic Canyon is globally high. However, such comparisons are invariably confounded by several factors including differences between studies in the range and area of habitats surveyed, and the types of sampling gear employed.

Comparisons of epibenthic diversity between studies are further complicated by the history and intensity of human impacts, such as demersal fishing. A number of reviews highlight the fact that demersal fishing gears such as beam-trawls, otter trawls and dredges modify benthic habitats and fauna (Dayton et al., 1995; Jennings and Kaiser, 1998; Thrush and Dayton, 2002). Typically demersal fishing gears dislodge attached epifauna and flatten existing topographic features. Significant mortalities of benthic species and modifications to benthic community structure are widely reported direct results of trawling and dredging impacts (Currie and Parry, 1996; Jennings and Kaiser, 1998). The time scales over which fishery induced community changes develop are also likely to vary with depth, as many deepwater species have life-history characteristics that differ from shelf species (i.e. increased longevity, slower growth rates and late maturation). Demersal trawling has been ongoing in the offshore waters of southeastern Australia for almost 100 years, and it is estimated that at least 65% of the upper slope and a large portion of the mid slope have been trawled in recent years (Wayte et al., 2006). While the cumulative effects of historical trawling impacts on the epibenthic assemblages of Bonney and du Couedic Canyon are unknown, it appears that trawling is becoming increasingly concentrated in Australia’s southern canyons, due in part to improvements in navigational technologies (Williams et al., 2009). We therefore cannot discount the possibility that patterns in

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epifauna distribution observed in this study reflect to some degree the persistent effects of historical fishing.

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Currie, D.R., 2008, The role of submarine canyons in upwelling, sediment transport, and productivity hotspots off the Bonney Coast and Kangaroo Island, South Australia. Southern Surveyor Voyage Summary SS02/2008., Report to the Marine National Facility: Hobart, CSIRO Marine and Atmospheric Research, p. 19. Currie, D.R., and Parry, G.D., 1996, Effects of scallop dredging on a soft sediment community: a large-scale environmental study: Marine Ecology Progress Series, v. 134, p. 131-150. Dayton, P.K., Thrush, S.F., Agardy, M.T., and Hofman, R.J., 1995, Environmental effects of marine fishing: Aquatic Conservation-Marine and Freshwater Ecosystems, v. 5, p. 205- 232. Diaz, M.C., and Klaus, R., 2001, Sponges: an essential component of Caribbean coral reefs: Bulletin of Marine Science, v. 69, p. 535-546. Fine, R.A., 1993, Circulation of Antarctic intermediate water in the South Indian Ocean: Deep- Sea Research Part I-Oceanographic Research Papers, v. 40, p. 2021-2042. Genin, A., 2004, Bio-physical coupling in the formation of zooplankton and fish aggregations over abrupt topographies: Journal of Marine Systems, v. 50, p. 3-20. Greene, C.H., Wiebe, P.H., Burczynski, J., and Youngbluth, M.J., 1988, Acoustical detection of high density krill demersal layers in the submarine canyons of Georges Bank: Science, v. 241, p. 359-361. Griffin, D.A., Thompson, P.A., Bax, N.J., Bradford, R.W., and Hallegraeff, G.M., 1997, The 1995 mass mortality of pilchard: No role found for physical or biological oceanographic factors in Australia: Marine and Freshwater Research, v. 48, p. 27-42. Hall, S.J., 1994, Physical disturbance and marine benthic communities: life in unconsolidated sediments: Oceanography and Marine Biology, v. 32, p. 179-239. Hill, P.J., De Deckker, P., and Exon, N.F., 2005, Geomorphology and evolution of the gigantic Murray canyons on the Australian southern margin: Australian Journal of Earth Sciences, v. 52, p. 117-136. Jennings, S., and Kaiser, M.J., 1998, The effects of fishing on marine ecosystems: Advances in Marine Biology, v. 34, p. 203-314. Johnson, N.A., Campbell, J.W., Moorre, T.S., Rex, M.A., Etter, R.J., McClain, C.R., and Dowell, M.D., 2007, The relationship between the standing stock of deep-sea macrobenthos and surface production in the western North Atlantic: Deep-Sea Research Part I- Oceanographic Research Papers, v. 54, p. 1350-1360. Koslow, J.A., Bulman, C.M., and Lyle, J.M., 1994, The mid-slope demersal fish community off southeastern Australia: Deep-Sea Research Part I-Oceanographic Research Papers, v. 41, p. 113-141.

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Koslow, J.A., Gowlett-Holmes, K., Lowry, J.K., O’Hara, T., Poore, G.C.B., and Williams, A., 2001, Seamount benthic macrofauna off southern Tasmania: community structure and impacts of trawling: Marine Ecology Progress Series, v. 213. Kroncke, I., Turkay, M., and Fiege, D., 2003, Macrofauna communities in the eastern Mediterranean deep sea: Marine Ecology, v. 24, p. 193-216. Lampitt, R.S., Raine, R.C.T., Billett, D.S.M., and Rice, A.L., 1995, Material supply to the European continental slope: A budget based on benthic oxygen demand and organic supply: Deep-Sea Research Part I-Oceanographic Research Papers, v. 42, p. 1865- 1880. Lennon, G.W., Bowers, D., Nunes, R.A., Scott, B.D., Ali, M., Boyle, J., Wenju, C., Herzfeld, M., Johansswon, G., Nield, S., Petrusevics, P., Suskin, A., and Wijffles, S.E.A., 1987, Gravity currents and the release of salt from and inverse estuary: Nature, v. 327, p. 695- 697. Lewis, R.K., 1981, Seasonal upwelling along the south-eastern coastline of South Australia: Australian Journal of Marine and Freshwater Research, v. 32, p. 843-854. Louzao, M., Anadon, N., Arrontes, J., Alvarez-Claudio, C., Fuente, D.M., Ocharan, F., Anadon, A., and Acuna, J.L., 2010, Historical macrobenthic community assemblages in the Aviles Canyon, N Iberian Shelf: baseline biodiversity information for a marine protected area: Journal of Marine Systems, v. 80, p. 47-56. Macquart-Moulin, C., and Patriti, G., 1996, Accumulation of migratory micronekton crustaceans over the upper slope and submarine canyons of the northwestern Mediterranean: Deep- Sea Research Part I-Oceanographic Research Papers, v. 43, p. 579-601. McClatchie, S., Middleton, J.F., and Ward, T.M., 2006, Water mass analysis and alongshore variation in upwelling intensity in the eastern Great Australian Bight: Journal of Geophysical Research, v. 111, p. doi: 10.1029/2004JC002699. Middleton, J.F., and Bye, J.A.T., 2007, A review of the shelf-slope circulation along Australia's southern shelves: Cape Leeuwin to Portland: Progress in Oceanography, v. 75, p. 1-41. Middleton, J.F., and Platov, G., 2003, The mean summertime circulation along Australia's southern shelves: A numerical study: Journal of Physical Oceanography, v. 33, p. 2270- 2287. Nieblas, A.E., Sloyan, B.M., Hobday, A.J., Coleman, R., and Richardson, A.J., 2009, Variability of biological production in low wind-forced regional upwelling systems: A case study off southeastern Australia: Limnology and Oceanography, v. 54, p. 1548-1558. Sanchez, F., Serrano, A., Parra, S., Ballesteros, M., and Cartes, J.E., 2008, Habitat characteristics as a determinant of the structure and spatial distribution of epibenthic

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and demersal communities of Le Danois Bank, Cantabrian Sea, N. Spain: Journal of Marine Systems, v. 72, p. 64-86. Schahinger, R.B., 1987, Structure of coastal upwelling events observed off the southeast coast of South Australia during February1983 - April 1984: Australian Journal of Marine and Freshwater Research, v. 38, p. 439-459. Smetacek, V., 1984, The supply of food to the benthos, in Fasham, M.J.R., ed., Flows of energy and materials in marine ecosystems: theory and practice: New York, Published in cooperation with NATO Scientific Affairs Division by Plenum Press, p. 517-547. Smith, S.V., and Veeh, H.H., 1987, Mass balance of biochemically active materials (C, N, P) in a hypersaline gulf: Estuarine, Coastal and Shelf Science, v. 29, p. 195-215. Snelgrove, P.V.R., and Butman, C.A., 1994, Animal sediment relationships revisited: cause versus effect, Oceanography and Marine Biology, Vol 32, Volume 32: Oceanography and Marine Biology, p. 111-177. Thouzeau, G., Robert, G., and Ugarte, R., 1991, Faunal assemblages of benthic megainvertebrates inhabiting sea scallop grounds from the eastern Georges Bank, in relation to environmental factors: Marine Ecology Progress Series, v. 74, p. 61-82. Thrush, S.F., and Dayton, P.K., 2002, Disturbance to marine benthic habitats by trawling and dredging: Implications for marine biodiversity: Annual Review of Ecology and Systematics, v. 33, p. 449-473. Ward, T.M., McLeay, L.J., Dimmlich, W.F., Rogers, P.J., McClatchie, S., Matthews, R., Kampf, J., and Van Ruth, P.D., 2006, Pelagic ecology of a northern boundary current system: effects of upwelling on the productivity and distribution of sardine Sardinops sagax, anchovy Engraulis australis and southern bluefin tuna Thunnus maccoyii in the Great Australian Bight: Fisheries Oceanography, v. 15, p. 191-207. Wayte, S., Dowdney, J., Williams, A., Bulman, C., Sporcic, M., Fuller, M., and Smith, A., 2006, Ecological Risk Assessment for the Effects of Fishing: Otter Trawl Component of the Southern and Eastern Scalefish and Shark Fishery Report, Report for the Australian Fisheries Management Authority: Canberra, p. 268. Williams, A., Bax, N.J., Kloser, R.J., Althaus, F., Barker, B., and Keith, G., 2009, Australia's deep-water reserve network: implications of false homogeneity for classifying abiotic surrogates of biodiversity: ICES Journal of Marine Science, v. 66, p. 214-224. Williams, A., Koslow, J.A., and Last, P.R., 2001, Diversity, density and community structure of the demersal fish fauna of the continental slope off western Australia, 20 to 35 degrees S: Marine Ecology-Progress Series, v. 212, p. 247-263.

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AKNOWLEDGEMENTS This work could not have been undertaken without the assistance of a highly skilled group of mariners. In particular, we would like to thank the Captain (Ian Taylor) and the crew of the RV Southern Surveyor for their support throughout the voyage. Thanks are also due the following Marine National Facility staff for their contributions to a highly successful voyage; Fred Stein (Director), Don McKenzie (Voyage Manager), Steve Thomas (Electronics), Bob Beattie (Computing), Mark Rayner (Hydrochemistry) and Anne Kennedy (Swath Mapping). We are also very much indebted to the following voyage scientists for their help on this research project; Graham Hooper and Mike Steer (SARDI), James Paterson (Flinders University), Wayne Rumball (SAM) and Ruan Gannon (University of Adelaide). Thanks also to Neil Chigwidden and the many support staff at SARDI who helped develop and mobilize the sampling equipment. This project was jointly funded by the South Australian Government (through the Wildlife Conservation Fund and the Marine Innovation South Australia initiative) and the Australian Federal Government.

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Figure 1. Map showing the locations of Bonney and du Couedic canyons (unfilled rectangles) on the south-east continental margin. Contour lines presented follow 100 m depth intervals.

136° E 137° E 138° E 139° E 140° E 141° E

South Australia Spencer Gulf

Adelaide 35° S Gulf 35° S St Vincent

Kangaroo Island 36° S 36° S

Lacepede du Couedic Shelf Canyon Victoria

37° S 37° S

Bonney Coast

Bonney Canyon Abyssal Plain 38° S 38° S

025 50 100 150 km

136° E 137° E 138° E 139° E 140° E 141° E

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Figure 2. Bathymetric map of du Couedic Canyon showing the locations of 5 survey sites sampled for macro-epibenthos, sediments and near-bed water properties. Labels denote location relative to the main canyon axis (DCC = du Couedic Centre) and sampling depth in metres (100, 200, 500, 1000, 1500).

136°20'E 136°30'E

Depth (m) 100

125

DCC_100 150

200

1500 36°20'S 36°20'S

DCC_200

DCC_500

36°30'S 36°30'S

DCC_1000

DCC_1500

02 4 8 km

136°20'E 136°30'E

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Figure 3. Bathymetric map of Bonney Canyon showing the locations of 6 survey sites sampled for macro-epibenthos, sediments and near-bed water properties. Labels denote location relative to the main canyon axis (BC = Bonney Centre) and sampling depth in metres (100, 200, 500, 1000, 1500, 2000).

139°20'E 139°30'E 139°40'E

Depth (m) 70

BC_100 300 37°30'S 37°30'S

800

1200

2100

BC_200

BC_500

37°40'S 37°40'S BC_1000

BC_1500

BC_2000

37°50'S 37°50'S

0 2.5 5 10 km

139°20'E 139°30'E 139°40'E

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Figure 4. Plots of (a) mean wet weight, (b) total number of species of each major phylum collected during the survey, and (c) total number of sites (out of 11) at which specimens belonging to each major phylum were collected. Values for each variable are shown as percentages above each bar.

(a) 200

83.94% 150

100

50 Biomass (kg/ha) 4.06% 7.32% 0.03% 0.03% <0.01% 1.07% 0.05% 1.98% <0.01% 1.51% 0

(b) 100 47.38% 80

60

40

Species(N) 11.96% 13.04% 20 8.15% 3.80% 3.80% 4.89% 4.35% 0.54% 1.63% 0.54% 0

(c) 12 100.00% 10 81.82% 81.82% 81.82% 72.73% 8 72.73% 72.73%

6 36.36% 4 27.27% Stations (N) 2 9.09% 9.09% 0 a a t a a d a t a d o a ta ia m a a ra a id o r r c e d l p p o a a e s rt fe r e o io z rd d u e ri o n r h o i d ll h n h c ry o n o o m o c rt h C in e P o A ra B C h M r A B c N U E

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Figure 5. Total (a) number of species, and (b) wet weights of macro-epibenthos collected during sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons.

(a)

100

80 du Couedic Bonney

60

40 Species(N)

20

0 (b)

10000

1000

100

10

Biomass(kg/ha) 1

0 100 200 500 1000 1500 2000

Depth (m)

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Figure 6. Dendrogram and non-metric MDS ordination of epifaunal community structure at 11 depth- stratified sampling stations at Bonney and du Couedic canyons. Sampling depths are represented by different symbols (upright triangle = 100 m, diamond = 200 m, circle = 500 m, inverted triangle = 1000 m, square = 1500 m, plus = 2000 m) and canyon locations by different shades of fill (black = Bonney, white = du Couedic). Three station groupings are identified at the 5% Bray-Curtis similarity level (dotted line): shelf (100-200 m), upper slope (500 m) and mid-slope (1000-2000 m).

(a) 0

20

40

60 Similarity 80

100

BC_100 BC_200 BC_500 BC_1000 BC_1500 BC_2000 DCC_200 DCC_100 DCC_500 DCC_1500 DCC_1000 Sampling stations

(b)

Stress = 0.02 BC_1000 BC_100

BC_200 DCC_100 DCC_1000 DCC_1500 DCC_200 Shelf

Mid Slope BC_500

BC_1500 Upper Slope

DCC_500

BC_2000

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Table 1. Summary of environmental factors at 11 epibenthic sampling sites at Bonney and du Couedic canyons.

Station Latitude Longitude Depth Temperature Salinity Chlorophyll Oxygen Sediment Sediment Sediment Sediment (ºS) (ºE) (m) (ºC) (µg/l) (µM/l) size sorting carbon nitrogen (%<63µm) (Phi) (% organic) (%) DCC_100 36.2825 136.5362 100 11.08 34.98 15.54 244.15 2.36 1.10 0.23 0.06 DCC_200 36.3916 136.4859 200 11.21 35.00 13.16 247.14 2.79 1.30 0.23 0.06 DCC_500 36.4561 136.4585 500 8.98 34.64 10.34 249.07 2.65 1.06 0.41 0.01 DCC_1000 36.5358 136.4092 1000 5.09 34.39 11.40 197.79 26.40 1.27 <0.01 0.13 DCC_1500 36.5811 136.4128 1500 2.87 34.54 11.33 172.24 43.66 1.44 <0.01 0.10

BC_100 37.5019 139.6152 100 10.69 34.91 13.87 252.69 0.97 1.33 0.06 0.06 BC_200 37.6010 139.5762 200 9.65 34.74 10.33 249.91 2.04 0.96 0.05 0.01 BC_500 37.6420 139.5383 500 8.92 34.63 10.34 249.67 5.93 0.83 0.26 0.06 BC_1000 37.6936 139.4830 1000 4.37 34.39 10.05 194.15 54.43 1.81 1.05 0.16 BC_1500 37.7326 139.4375 1500 2.66 34.60 10.73 170.68 56.61 1.68 1.07 0.16 BC_2000 37.8037 139.3922 2000 ------

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Table 2. Spearman’s rank correlation coefficients between epifaunal richness and biomass and adjacent environmental conditions at 11 depth-stratified sampling stations at Bonney (B) and du Couedic (dC) Cannon. Significant positive (+) and negative (-) correlations are denoted at the ** 1% and * 5% level.

Sediment Sediment Sediment Depth Temperature Salinity Chlorophyll Oxygen % Mud Richness sorting carbon nitrogen Depth ...... Temperature -B**,-dC* ...... Salinity -B* +B*,+dC* ...... Chlorophyll – – – ...... Oxygen -B** +B** +B* – ...... % Mud +B**,+dC* -B** -B* – -B** . . . . . Sediment sorting – – – – – – . . . . Sediment carbon +B*,+dC** -B*,-dC* – – -B* +B*,+dC* – . . . Sediment nitrogen – – – – – – – +B** . . Richness – +dC* +dC** – – – – – – . Biomass -dC** +dC* – – – -dC* – -dC** – +B*

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Table 3. Mean biomass (kg per ha ± s.e.) of macro-epibenthic species in three station groups identified from MDS classification. Species listed were identified from SIMPER analyses as contributing ≥5% to the similarity within and dissimilarity between regional groupings. Those species indicative of each regional grouping (contributing ≥10% to the total similarity within a group) are highlighted in bold. Species are ranked in order of decreasing biomass across all station groupings.

Phylum Scientific Name Shelf (n = 4) Upper Slope (n = 2) Mid Slope (n = 5) Porifera Spirophorid sp. 4.46 ± 4.46 Porifera Spheciospongia papillosa 27.46 ± 24.26 Porifera Spheciospongia sp. 38.30 ± 20.86 Mollusca Pyxipoma weldii 37.84 ± 32.71 Porifera Demosponge sp. 14 16.09 ± 10.09 1.20 ± 1.20 Porifera Chalinid sp. 16.51 ± 8.53 Porifera Jaspis lutea 5.59 ± 3.07 Cnidaria Clathrozoon wilsoni 4.14 ± 1.04 Echinodermata Astropecten sp. 1 0.29 ± 0.14 Echinodermata Holothuroid sp. 2 0.23 ± 0.14 Porifera Polymastia sp. 2 24.81 ± 18.82 Echinodermata Astropecten sp. 3 0.16 ± 0.10 Echinodermata Brissus sp. 0.41 ± 0.01 Echinodermata Astropecten sp. 4 0.12 ± 0.08 Mollusca Lucinoma galatheae 0.02 ± 0.01 Urochordata Thalacian sp. 1 0.02 ± 0.01

SARDI Aquatic Sciences Report – Page 30 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Appendix 1. Taxonomic and functional classification of 184 macro-epibenthic species collected from sled tows at 11 depth-stratified sampling stations at Bonney and du Couedic canyons. All species codes given here refer to material lodged in the South Australian Museum.

Phylum Class ScientificName CommonName Diet Mobility Code Porifera Calcarea Clathrina sp. Sponge Suspension Sessile I248 Porifera Demospongiae Jaspis stellifera Sponge Suspension Sessile I056 Porifera Demospongiae Fasciospongia sp. Sponge Suspension Sessile I094 Porifera Demospongiae Chalinid sp. Sponge Suspension Sessile I095 Porifera Demospongiae Axinella sp. 1 Sponge Suspension Sessile I096 Porifera Demospongiae Spheciospongia sp. Sponge Suspension Sessile I097 Porifera Demospongiae Leiosella sp. Sponge Suspension Sessile I098 Porifera Demospongiae Jaspis lutea Sponge Suspension Sessile I099 Porifera Demospongiae Dictyoceratid sp. 1 Sponge Suspension Sessile I100 Porifera Demospongiae Dictyoceratid sp. 2 Sponge Suspension Sessile I101 Porifera Demospongiae Geodiid sp. Sponge Suspension Sessile I102 Porifera Demospongiae Dendroceratid sp. 1 Sponge Suspension Sessile I103 Porifera Demospongiae Hemiasterella sp. 1 Sponge Suspension Sessile I105 Porifera Demospongiae Stelletta sp. Sponge Suspension Sessile I106 Porifera Demospongiae Cymbastela sp. Sponge Suspension Sessile I107 Porifera Demospongiae Dendroceratid sp. 2 Sponge Suspension Sessile I108 Porifera Demospongiae Spheciospongia papillosa Sponge Suspension Sessile I109 Porifera Demospongiae Suberitid sp. Sponge Suspension Sessile I110 Porifera Demospongiae Polymastia sp. 1 Sponge Suspension Sessile I113 Porifera Demospongiae Oceanapia sp. 1 Sponge Suspension Sessile I114 Porifera Demospongiae Axinella sp. 2 Sponge Suspension Sessile I115 Porifera Demospongiae Phakellia sp. Sponge Suspension Sessile I116 Porifera Demospongiae Hippospongia sp. Sponge Suspension Sessile I117 Porifera Demospongiae Demosponge sp. 1 Sponge Suspension Sessile I119 Porifera Demospongiae Demosponge sp. 2 Sponge Suspension Sessile I120 Porifera Demospongiae Dictyoceratid sp. 3 Sponge Suspension Sessile I121 Porifera Demospongiae Taonura sp. Sponge Suspension Sessile I122 Porifera Demospongiae Cribrochalina sp. 1 Sponge Suspension Sessile I123 Porifera Demospongiae Siphonochalina sp. Sponge Suspension Sessile I124 Porifera Demospongiae Raspailia sp. Sponge Suspension Sessile I125 Porifera Demospongiae Dictyoceratid sp. 4 Sponge Suspension Sessile I126 Porifera Demospongiae Rhizaxinella sp. Sponge Suspension Sessile I127 Porifera Demospongiae Oceanapia sp. 2 Sponge Suspension Sessile I128 Porifera Demospongiae Dictyoceratid sp. 5 Sponge Suspension Sessile I129 Porifera Demospongiae Dictyoceratid sp. 6 Sponge Suspension Sessile I130 Porifera Demospongiae Strongylacidon sp. Sponge Suspension Sessile I132 Porifera Demospongiae Polymastia sp. 2 Sponge Suspension Sessile I133 Porifera Demospongiae Demosponge sp. 3 Sponge Suspension Sessile I134 Porifera Demospongiae Dendroceratid sp. 3 Sponge Suspension Sessile I136 Porifera Demospongiae Demosponge sp. 4 Sponge Suspension Sessile I137 Porifera Demospongiae Demosponge sp. 5 Sponge Suspension Sessile I138 Porifera Demospongiae Demosponge sp. 6 Sponge Suspension Sessile I139 Porifera Demospongiae Demosponge sp. 7 Sponge Suspension Sessile I140 Porifera Demospongiae Ircinia sp. 1 Sponge Suspension Sessile I141 Porifera Demospongiae Cribrochalina sp. 2 Sponge Suspension Sessile I145 Porifera Demospongiae Demosponge sp. 8 Sponge Suspension Sessile I147 Porifera Demospongiae Demosponge sp. 9 Sponge Suspension Sessile I148 Porifera Demospongiae Demosponge sp. 10 Sponge Suspension Sessile I149 Porifera Demospongiae Oceanapia sp. 3 Sponge Suspension Sessile I150 Porifera Demospongiae Dictyoceratid sp. 7 Sponge Suspension Sessile I151

SARDI Aquatic Sciences Report – Page 31 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Phylum Class ScientificName CommonName Diet Mobility Code Porifera Demospongiae Dictyoceratid sp. 8 Sponge Suspension Sessile I152 Porifera Demospongiae Dictyoceratid sp. 9 Sponge Suspension Sessile I154 Porifera Demospongiae Demosponge sp. 11 Sponge Suspension Sessile I155 Porifera Demospongiae Demosponge sp. 12 Sponge Suspension Sessile I156 Porifera Demospongiae Demosponge sp. 13 Sponge Suspension Sessile I157 Porifera Demospongiae Dictyonella sp. Sponge Suspension Sessile I159 Porifera Demospongiae Demosponge sp. 14 Sponge Suspension Sessile I161 Porifera Demospongiae Dictyoceratid sp. 10 Sponge Suspension Sessile I163 Porifera Demospongiae Psammocinia sp. Sponge Suspension Sessile I165 Porifera Demospongiae Demosponge sp. 15 Sponge Suspension Sessile I174 Porifera Demospongiae Demosponge sp. 16 Sponge Suspension Sessile I175 Porifera Demospongiae Demosponge sp. 17 Sponge Suspension Sessile I182 Porifera Demospongiae Demosponge sp. 18 Sponge Suspension Sessile I185 Porifera Demospongiae Demosponge sp. 19 Sponge Suspension Sessile I197 Porifera Demospongiae Demosponge sp. 20 Sponge Suspension Sessile I203 Porifera Demospongiae Demosponge sp. 21 Sponge Suspension Sessile I204 Porifera Demospongiae Demosponge sp. 22 Sponge Suspension Sessile I205 Porifera Demospongiae Demosponge sp. 23 Sponge Suspension Sessile I212 Porifera Demospongiae Demosponge sp. 24 Sponge Suspension Sessile I233 Porifera Demospongiae Spirophorid sp. Sponge Suspension Sessile I234 Porifera Demospongiae Demosponge sp. 25 Sponge Suspension Sessile I235 Porifera Demospongiae Demosponge sp. 26 Sponge Suspension Sessile I236 Porifera Demospongiae Demosponge sp. 27 Sponge Suspension Sessile I237 Porifera Demospongiae Dysidea sp. Sponge Suspension Sessile I238 Porifera Demospongiae Demosponge sp. 28 Sponge Suspension Sessile I239 Porifera Demospongiae Demosponge sp. 29 Sponge Suspension Sessile I241 Porifera Demospongiae Demosponge sp. 30 Sponge Suspension Sessile I242 Porifera Demospongiae Oceanapia sp. 4 Sponge Suspension Sessile I243 Porifera Demospongiae Ancorinid sp. Sponge Suspension Sessile I244 Porifera Demospongiae Demosponge sp. 31 Sponge Suspension Sessile I245 Porifera Demospongiae Clathria sp. 1 Sponge Suspension Sessile I247 Porifera Demospongiae Echinodictyum sp. Sponge Suspension Sessile I255 Porifera Demospongiae Ircinia sp. 2 Sponge Suspension Sessile I256 Porifera Demospongiae Halichondrid sp. Sponge Suspension Sessile I257 Porifera Demospongiae Aka sp. Sponge Suspension Sessile I258 Porifera Demospongiae Poecilosclerid sp. Sponge Suspension Sessile I259 Porifera Hexactinellida Hexactinellid sp. Sponge Suspension Sessile I221 Cnidaria Anthozoa Umbellulifera sp. Soft coral Suspension Sessile I090 Cnidaria Anthozoa Pteronisis sp. 1 Sea fan Suspension Sessile I092 Cnidaria Anthozoa Dendronephthea sp. Soft coral Suspension Sessile I111 Cnidaria Anthozoa Nephtheid sp. Soft coral Suspension Sessile I153 Cnidaria Anthozoa Isidid sp. 1 Sea fan Suspension Sessile I162 Cnidaria Anthozoa Oculinid sp. Coral Suspension Sessile I171 Cnidaria Anthozoa Actiniarid sp. 1 Anemone Suspension Sessile I172 Cnidaria Anthozoa Scleractinid sp. Coral Suspension Sessile I181 Cnidaria Anthozoa Caryophyllia planilamellata Coral Suspension Sessile I199 Cnidaria Anthozoa Actiniarid sp. 2 Anemone Suspension Sessile I201 Cnidaria Anthozoa Alcyonacid sp. 1 Sea fan Suspension Sessile I206 Cnidaria Anthozoa Umbellula sp. Sea pen Suspension Sessile I226 Cnidaria Anthozoa Isidid sp. 2 Sea whip Suspension Sessile I240 Cnidaria Anthozoa Alcyonacid sp. 2 Sea fan Suspension Sessile I251 Cnidaria Anthozoa Desmophyllum dianthus Coral Suspension Sessile I292 Cnidaria Anthozoa Alcyonacid sp. 3 Sea fan Suspension Sessile I293

SARDI Aquatic Sciences Report – Page 32 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Phylum Class ScientificName CommonName Diet Mobility Code Cnidaria Hydrozoa Sertularia sp. Hydroid Suspension Sessile I091 Cnidaria Hydrozoa Clathrozoon wilsoni Hydroid Suspension Sessile I093 Cnidaria Hydrozoa Hydroid sp. 1 Hydroid Suspension Sessile I169 Cnidaria Hydrozoa Stylaster sp. Hydocoral Suspension Sessile I186 Cnidaria Hydrozoa Hydroid sp. 2 Hydroid Suspension Sessile I193 Cnidaria Hydrozoa Hydroid sp. 3 Hydroid Suspension Sessile I208 Nemertea Enopla Nermertean sp. Ribbon worm Scavenger Mobile I230 Annelida Polychaeta Serpulid sp. 1 Worm Suspension Sessile I166 Annelida Polychaeta Chaetopterid sp. Worm Deposit Sessile I167 Annelida Polychaeta Maldanid sp. 2 Worm Deposit Sessile I176 Annelida Polychaeta Maldanid sp. 1 Worm Deposit Sessile I177 Annelida Polychaeta Scalibregma sp. Worm Deposit Mobile I209 Annelida Polychaeta Serpulid sp. 2 Worm Suspension Sessile I294 Annelida Polychaeta Serpulid sp. 3 Worm Suspension Sessile I295 Arthropoda Malacostraca Brucerolis victoriensis Isopod Deposit Mobile I009 Arthropoda Malacostraca Sphaeromid sp. Pill bug Scavenger Mobile I158 Arthropoda Malacostraca Pagurid sp. Crab Scavenger Mobile I189 Arthropoda Malacostraca Majid sp. Spider crab Scavenger Mobile I195 Arthropoda Malacostraca Galathea sp. Squat lobster Scavenger Mobile I196 Arthropoda Malacostraca Ebalia sp. Crab Scavenger Mobile I211 Arthropoda Pycnogonida Colossendeis tasmanica Sea spider Predator Mobile I007 Mollusca Bivalvia Lucinoma galatheae Shell Suspension Sessile I170 Mollusca Bivalvia Anomiid sp. Shell Suspension Sessile I200 Mollusca Bivalvia Semipallium aktinos Shell Suspension Sessile I246 Mollusca Bivalvia Mytilid sp. Shell Suspension Sessile I254 Mollusca Bivalvia Neotrigonia sp. Shell Suspension Sessile I290 Mollusca Gastropoda Trochid sp. Shell Predator Mobile I190 Mollusca Gastropoda Agnewia sp. Shell Predator Mobile I228 Mollusca Gastropoda Pyxipoma weldii Shell Predator Mobile I272 Mollusca Cephalopoda Octopoteuthis sp. Squid beak Predator Mobile I229 Brachiopoda Inarticulata Brachiopod sp. Lamp shell Suspension Sessile I223 Bryozoa Gymnolaemata Adeona spp. Lace coral Suspension Sessile I112 Bryozoa Gymnolaemata Bryozoan sp. 1 Lace coral Suspension Sessile I131 Bryozoa Gymnolaemata Triphyllozoon sp. Lace coral Suspension Sessile I135 Bryozoa Gymnolaemata Orthoscuticella spp. Lace coral Suspension Sessile I142 Bryozoa Gymnolaemata Bryozoan sp. 2 Lace coral Suspension Sessile I146 Bryozoa Gymnolaemata Bryozoan sp. 3 Lace coral Suspension Sessile I168 Bryozoa Gymnolaemata Bryozoan sp. 4 Lace coral Suspension Sessile I173 Bryozoa Gymnolaemata Bryozoan sp. 5 Lace coral Suspension Sessile I187 Bryozoa Gymnolaemata Iodictyum phoeniceum Lace coral Suspension Sessile I191 Bryozoa Gymnolaemata Bryozoan sp. 6 Lace coral Suspension Sessile I192 Bryozoa Gymnolaemata Bryozoan sp. 7 Lace coral Suspension Sessile I207 Bryozoa Gymnolaemata Bryozoan sp. 8 Lace coral Suspension Sessile I214 Bryozoa Gymnolaemata Bryozoan sp. 9 Lace coral Suspension Sessile I253 Bryozoa Gymnolaemata Phidoloporid sp. 1 Lace coral Suspension Sessile I291 Bryozoa Stenolaemata Hornera robusta Lace coral Suspension Sessile I164 Echinodermata Asteroidea Astropecten sp. 1 Sea star Scavenger Mobile I008 Echinodermata Asteroidea Brisingid sp. Sea star Scavenger Mobile I023 Echinodermata Asteroidea Astropecten preissei Sea star Scavenger Mobile I188 Echinodermata Asteroidea Asteroid sp. 1 Sea star Scavenger Mobile I215 Echinodermata Asteroidea Astropecten sp. 2 Sea star Scavenger Mobile I216 Echinodermata Asteroidea Astropecten sp. 3 Sea star Scavenger Mobile I217 Echinodermata Asteroidea Astropecten sp. 4 Sea star Scavenger Mobile I218

SARDI Aquatic Sciences Report – Page 33 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Phylum Class ScientificName CommonName Diet Mobility Code Echinodermata Asteroidea Asteroid sp. 2 Sea star Scavenger Mobile I289 Echinodermata Crinoidea Ptilometra sp. Feather star Suspension Mobile I232 Echinodermata Echinoidea Goniocidaris tubaria Urchin Grazer Mobile I104 Echinodermata Echinoidea Brissus sp. Urchin Deposit Mobile I194 Echinodermata Holothuroidea Holothuroid sp. 1 Sea cucumber Deposit Mobile I179 Echinodermata Holothuroidea Trochodota sp. Sea cucumber Deposit Mobile I180 Echinodermata Holothuroidea Holothuroid sp. 2 Sea cucumber Deposit Mobile I220 Echinodermata Ophiuroidea Ophuiroid sp. 1 Brittle star Scavenger Mobile I143 Echinodermata Ophiuroidea Ophuiroid sp. 2 Brittle star Scavenger Mobile I144 Echinodermata Ophiuroidea Ophuiroid sp. 3 Brittle star Scavenger Mobile I183 Echinodermata Ophiuroidea Ophiurid sp. 1 Brittle star Scavenger Mobile I184 Echinodermata Ophiuroidea Ophiothrix sp. Brittle star Scavenger Mobile I198 Echinodermata Ophiuroidea Ophiurid sp. 2 Brittle star Scavenger Mobile I210 Echinodermata Ophuiroidea Ophuiroid sp. 6 Brittle star Scavenger Mobile I219 Echinodermata Ophuiroidea Ophuiroid sp. 7 Brittle star Scavenger Mobile I224 Echinodermata Ophuiroidea Ophuiroid sp. 8 Brittle star Scavenger Mobile I225 Echinodermata Ophuiroidea Ophuiroid sp. 9 Brittle star Scavenger Mobile I227 Urochordata Thaliacea Thalacian sp. 1 Salp Suspension Planktonic I213 Urochordata Thaliacea Thalacian sp. 2 Salp Suspension Planktonic I222 Urochordata Ascidiacea Didemnid sp. 1 Sea squirt Suspension Sessile I160 Urochordata Ascidiacea Ascidian sp. 1 Sea squirt Suspension Sessile I202 Urochordata Ascidiacea Ascidian sp. 2 Sea squirt Suspension Sessile I249 Urochordata Ascidiacea Herdmania momus Sea squirt Suspension Sessile I250 Urochordata Ascidiacea Ascidian sp. 3 Sea squirt Suspension Sessile I252 Urochordata Ascidiacea Ascidian sp. 4 Sea squirt Suspension Sessile I260 Chordata Osteichthyes Helicolenus percoides Fish Predator Mobile F026 Chordata Osteichthyes Hoplichthys haswelli Fish Predator Mobile F028 Chordata Osteichthyes Morid sp. Fish Predator Mobile F041

SARDI Aquatic Sciences Report – Page 34 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Appendix 2. Photographic plates depicting 184 organisms collected in epibenthic sled tows at 11 depth- stratified sampling stations at Bonney and du Couedic canyons.

I248 – Clathrina sp. I097 – Spheciospongia sp. I102 – Geodiid sp.

I056 – Jaspis stellifera I098 – Leiosella sp. I103 – Dendroceratid sp. 1

I094 – Fasciospongia sp. I099 – Jaspis lutea I105 – Hemiasterella sp. 1

I095 – Chalinid sp. I100 – Dictyoceratid sp. 1 I106 – Stelletta sp.

I096 – Axinella sp. 1 I101 – Dictyoceratid sp. 2 I107 – Cymbastela sp.

SARDI Aquatic Sciences Report – Page 35 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I108 – Dendroceratid sp. 2 I115 – Axinella sp. 2 I121 – Dictyoceratid sp. 3

I109 – Spheciospongia papillosa I116– Phakellia sp. I122 – Taonura sp.

I110 – Suberitid sp. I117 – Hippospongia sp. I123 – Cribrochalina sp. 1

I113 – Polymastia sp. 1 I119 – Demosponge sp. 1 I124 – Siphonochalina sp.

I114 – Oceanapia sp. 1 I120 – Demosponge sp. 2 I125 – Raspailia sp.

SARDI Aquatic Sciences Report – Page 36 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I126 – Dictyoceratid sp. 4 I132 – Strongylacidon sp. I138 – Demosponge sp. 5

I127 – Rhizaxinella sp. I133 – Polymastia sp. 2 I139 – Demosponge sp. 6

I128 – Oceanapia sp. 2 I134 – Demosponge sp. 3 I140 – Demosponge sp. 7

I129 – Dictyoceratid sp. 5 I136 – Dendroceratid sp. 3 I141 – Ircinia sp. 1

I130 – Dictyoceratid sp. 6 I137 – Demosponge sp. 4 I145 – Cribrochalina sp. 2

SARDI Aquatic Sciences Report – Page 37 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I147 – Demosponge sp. 8 I152 – Dictyoceratid sp. I159 – Dictyonella sp.

I148 – Demosponge sp. 9 I154 – Dictyoceratid sp. 9 I161 – Demosponge sp. 14

I149 – Demosponge sp. 10 I155 – Demosponge sp. 11 I163 – Dictyoceratid sp. 10

I150 – Oceanapia sp. 3 I156 – Demosponge sp. 12 I165 – Psammocinia sp.

I151– Dictyoceratid sp. 7 I157 – Demosponge sp. 13 I174 – Demosponge sp. 15 (microscope image)

SARDI Aquatic Sciences Report – Page 38 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I175 – Demosponge sp. 16 (microscope image) I204 – Demosponge sp. 21 I235 – Demosponge sp. 25

I182 – Demosponge sp. 17 I205 – Demosponge sp. 22 I236 – Demosponge sp. 26

I185 – Demosponge sp. 18 I212 – Demosponge sp. 23 I237 – Demosponge sp. 27

I197 – Demosponge sp. 19 I133 – Demosponge sp. 24 I238 – Dysidea sp.

I203 – Demosponge sp. 20 I234 – Spirophorid sp. I239 – Demosponge sp. 28

SARDI Aquatic Sciences Report – Page 39 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I241 – Demosponge sp. 29 I247 – Clathria sp. 1 I259 - Poecilosclerid sp.

I242 – Demosponge sp. 30 I255 – Echinodictyum sp. I221 – Hexactinellid sp.

I243 – Oceanapia sp. I256 – Ircinia sp. 2 I090 – Umbellulifera sp.

I244 – Ancorinid sp. I257 – Halichondrid sp. I092 – Pteronisis sp. 1

I245 – Demosponge sp. 31 I258 – Aka sp. I111 – Dendronephthea sp.

SARDI Aquatic Sciences Report – Page 40 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I153 – Nephtheid sp. I201 – Actiniarid sp. 2 I292 – Desmophyllum dianthus

I162 – Isidid sp. 1 I206 – Alcyonacid sp. 1 I293 – Alcyonacid sp. 3

I171 – Oculinid sp. I226 – Umbellula sp. I091 – Sertularia sp.

I172 – Actiniarid sp. 1 I240 – Isidid sp. 2 I093 – Clathrozoon wilsoni

I199 – Caryophyllia planilamellata I251 – Alcyonacid sp. 2 I186 – Stylaster sp.

SARDI Aquatic Sciences Report – Page 41 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I193 – Hydroid sp. 2 I176 – Maldanid sp. 2 I009 – Brucerolis victoriensis

I208 – Hydroid sp. 3 I177 – Maldanid sp. 1 I158 – Sphaeromid sp.

I230 – Nermertean sp. I209 – Scalibregma sp. I189 – Pagurid sp.

I166 – Serpulid sp. 1 I294 – Serpulid sp. 2 I195 – Majid sp.

I167 – Chaetopterid sp. I295 – Serpulid sp. 3 I196 – Galathea sp.

SARDI Aquatic Sciences Report – Page 42 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I211 – Ebalia sp. I254 – Mytilid sp. I223 – Brachiopod sp.

I007 – Colossendeis tasmanica I290 – Neotrigonia sp. I112 – Adeona spp.

I170 – Lucinoma galatheae I190 – Trochid sp. I131 – Bryozoan sp. 1

I200 – Anomiid sp. I228 – Agnewia sp. I135 – Triphyllozoon sp.

I246 – Semipallium aktinos I229 – Octopoteuthis sp. (beak) I142 – Orthoscuticella spp.

SARDI Aquatic Sciences Report – Page 43 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I146 – Bryozoan sp. 2 I192 – Bryozoan sp. 6 I008 – Astropecten sp. 1

I168 – Bryozoan sp. 3 I207 – Bryozoan sp. 7 I023 – Brisingid sp.

I173 – Bryozoan sp. 4 I253 – Bryozoan sp. 9 I188 – Astropecten preissei

I187 – Bryozoan sp. 5 I291 – Phidoloporid sp. 1 I215 – Asteroid sp. 1

I191 – Iodictyum phoeniceum I164 – Hornera robusta I216 – Astropecten sp. 2

SARDI Aquatic Sciences Report – Page 44 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I217 – Astropecten sp. 3 I194 – Brissus sp. I144 – Ophuiroid sp. 2

I218 – Astropecten sp. 4 I179 – Holothuroid sp. 1 I183 – Ophuiroid sp. 3

I289 – Asteroid sp. 2 I180 – Trochodota sp. I184 – Ophiurid sp. 1

I232 – Ptilometra sp. I220 – Holothuroid sp. 2 I198 – Ophiothrix sp.

I104 – Goniocidaris tubaria I143 – Ophuiroid sp. 1 I210 – Ophiurid sp. 2

SARDI Aquatic Sciences Report – Page 45 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

I219 – Ophuiroid sp. 6 I222 – Thalacian sp. 2 I252 – Ascidian sp. 3

I224 – Ophuiroid sp. 7 I160 – Didemnid sp. 1 I260 – Ascidian sp. 4

I225 – Ophuiroid sp. 8 I202 – Ascidian sp. 1 F026 – Helicolenus percoides

I227 – Ophuiroid sp. 9 I249 – Ascidian sp. 2 F028 – Hoplichthys haswelli

I213 – Thalacian sp. 1 I250 – Herdmania momus F028 – Morid sp. Photos not available for: I181 – Scleractinid sp. I169 – Hydroid sp. 1, I272 – Pyxipoma weldii; I214 – Bryozoan sp. 8

SARDI Aquatic Sciences Report – Page 46 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Appendix 3. Summary list of species biomasses (kg per hectare) collected during sled tows at 11 depth- stratified sampling stations at Bonney and du Couedic canyons.

Station Species Code Weight Station Species Code Weight BC_100 Adeona spp. I112 0.080 BC_1000 Brisingid sp. I023 3.780 BC_100 Alcyonacid sp. 3 I293 0.060 BC_1000 Brucerolis victoriensis I009 0.020 BC_100 Ascidian sp. 2 I249 3.000 BC_1000 Bryozoan sp. 5 I187 0.020 BC_100 Asteroid sp. 2 I289 0.040 BC_1000 Demosponge sp. 17 I182 0.160 BC_100 Astropecten preissei I188 0.140 BC_1000 Demosponge sp. 18 I185 0.020 BC_100 Bryozoan sp. 6 I192 0.020 BC_1000 Holothuroid sp. 1 I179 4.340 BC_100 Chalinid sp. I095 2.040 BC_1000 Ophiurid sp. 1 I184 0.012 BC_100 Clathria sp. 1 I247 2.000 BC_1000 Ophuiroid sp. 1 I143 0.010 BC_100 Clathrozoon wilsoni I093 3.600 BC_1000 Ophuiroid sp. 3 I183 0.014 BC_100 Didemnid sp. 1 I160 0.500 BC_1000 Scleractinid sp. I181 0.560 BC_100 Hydroid sp. 2 I193 0.020 BC_1000 Stylaster sp. I186 0.020 BC_100 Iodictyum phoeniceum I191 0.020 BC_1000 Trochodota sp. I180 0.160 BC_100 Jaspis lutea I099 1.000 BC_1500 Asteroid sp. 1 I215 0.400 BC_100 Ophiurid sp. 2 I210 0.020 BC_1500 Astropecten sp. 1 I008 0.840 BC_100 Orthoscuticella spp. I142 0.200 BC_1500 Astropecten sp. 2 I216 0.860 BC_100 Pagurid sp. I189 0.200 BC_1500 Astropecten sp. 3 I217 0.420 BC_100 Polymastia sp. 2 I133 3.000 BC_1500 Astropecten sp. 4 I218 0.260 BC_100 Pyxipoma weldii I272 2.500 BC_1500 Brachiopod sp. I223 0.010 BC_100 Spheciospongia papillosa I109 2.000 BC_1500 Brucerolis victoriensis I009 0.020 BC_100 Spheciospongia sp. I097 2.000 BC_1500 Hexactinellid sp. I221 1.960 BC_100 Trochid sp. I190 0.040 BC_1500 Holothuroid sp. 2 I220 0.520 BC_200 Adeona spp. I112 0.080 BC_1500 Ophuiroid sp. 1 I143 0.008 BC_200 Alcyonacid sp. 3 I293 0.016 BC_1500 Ophuiroid sp. 6 I219 0.840 BC_200 Asteroid sp. 2 I289 0.040 BC_1500 Ophuiroid sp. 7 I224 0.014 BC_200 Chalinid sp. I095 1.520 BC_1500 Ophuiroid sp. 8 I225 0.012 BC_200 Clathrozoon wilsoni I093 1.800 BC_1500 Thalacian sp. 2 I222 0.100 BC_200 Demosponge sp. 11 I155 10.000 BC_2000 Agnewia sp. I228 0.080 BC_200 Didemnid sp. 1 I160 0.400 BC_2000 Asteroid sp. 1 I215 0.140 BC_200 Desmophyllum dianthus I292 0.200 BC_2000 Astropecten sp. 3 I217 0.400 BC_200 Goniocidaris tubaria I104 1.600 BC_2000 Astropecten sp. 4 I218 0.360 BC_200 Iodictyum phoeniceum I191 0.030 BC_2000 Colossendeis tasmanica I007 0.040 BC_200 Jaspis lutea I099 4.000 BC_2000 Holothuroid sp. 2 I220 0.620 BC_200 Ophiurid sp. 2 I210 0.020 BC_2000 Morid sp. F041 0.140 BC_200 Orthoscuticella spp. I142 0.300 BC_2000 Nermertean sp. I230 0.020 BC_200 Pagurid sp. I189 0.200 BC_2000 Octopoteuthis sp. I229 0.020 BC_200 Phidoloporid sp. 1 I291 0.060 BC_2000 Ophuiroid sp. 6 I219 0.220 BC_200 Polymastia sp. 2 I133 2.000 BC_2000 Ophuiroid sp. 9 I227 0.040 BC_200 Pyxipoma weldii I272 1.000 BC_2000 Thalacian sp. 1 I213 0.060 BC_200 Scalibregma sp. I209 0.040 BC_2000 Umbellula sp. I226 0.080 BC_200 Serpulid sp. 2 I294 0.200 DC_100 Adeona spp. I112 0.129 BC_200 Serpulid sp. 2 I294 0.240 DC_100 Aka sp. I258 3.225 BC_200 Serpulid sp. 3 I295 0.100 DC_100 Alcyonacid sp. 2 I251 10.793 BC_200 Spheciospongia papillosa I109 1.000 DC_100 Ancorinid sp. I244 8.643 BC_200 Spheciospongia sp. I097 3.000 DC_100 Ascidian sp. 2 I249 17.630 BC_500 Brissus sp. I194 0.400 DC_100 Ascidian sp. 3 I252 0.473 BC_500 Ebalia sp. I211 0.010 DC_100 Ascidian sp. 4 I260 1.505 BC_500 Desmophyllum dianthus I292 0.240 DC_100 Axinella sp. 2 I115 3.741 BC_500 Ophiurid sp. 2 I210 0.020 DC_100 Bryozoan sp. 8 I214 0.430 BC_500 Phidoloporid sp. 1 I291 0.060 DC_100 Bryozoan sp. 9 I253 0.086 BC_500 Scalibregma sp. I209 0.040 DC_100 Chalinid sp. I095 32.981 BC_1000 Astropecten sp. 1 I008 0.168 DC_100 Clathria sp. 1 I247 3.311

SARDI Aquatic Sciences Report – Page 47 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Station Species Code Weight Station Species Code Weight DC_100 Clathrina sp. I248 0.387 DC_200 Cribrochalina sp. 1 I123 2.769 DC_100 Clathrozoon wilsoni I093 6.794 DC_200 Cribrochalina sp. 2 I145 5.152 DC_100 Demosponge sp. 13 I157 6.235 DC_200 Cymbastela sp. I107 2.834 DC_100 Demosponge sp. 14 I161 41.839 DC_200 Demosponge sp. 1 I119 13.975 DC_100 Demosponge sp. 2 I120 4.214 DC_200 Demosponge sp. 10 I149 0.966 DC_100 Demosponge sp. 24 I233 117.992 DC_200 Demosponge sp. 11 I155 25.502 DC_100 Demosponge sp. 25 I235 5.117 DC_200 Demosponge sp. 12 I156 29.817 DC_100 Demosponge sp. 26 I236 68.327 DC_200 Demosponge sp. 13 I157 9.660 DC_100 Demosponge sp. 27 I237 5.504 DC_200 Demosponge sp. 14 I161 22.540 DC_100 Demosponge sp. 28 I239 129.559 DC_200 Demosponge sp. 2 I120 5.023 DC_100 Demosponge sp. 29 I241 14.276 DC_200 Demosponge sp. 3 I134 0.515 DC_100 Demosponge sp. 30 I242 3.483 DC_200 Demosponge sp. 4 I137 0.193 DC_100 Demosponge sp. 31 I245 75.035 DC_200 Demosponge sp. 5 I138 0.580 DC_100 Dictyoceratid sp. 10 I163 15.824 DC_200 Demosponge sp. 6 I139 0.064 DC_100 Didemnid sp. 1 I160 0.387 DC_200 Demosponge sp. 7 I140 0.902 DC_100 Dysidea sp. I238 0.430 DC_200 Demosponge sp. 8 I147 1.932 DC_100 Echinodictyum sp. I255 5.891 DC_200 Demosponge sp. 9 I148 3.220 DC_100 Geodiid sp. I102 82.689 DC_200 Dendroceratid sp. 1 I103 0.258 DC_100 Goniocidaris tubaria I104 8.729 DC_200 Dendroceratid sp. 2 I108 4.637 DC_100 Halichondrid sp. I257 0.645 DC_200 Dendroceratid sp. 3 I136 1.030 DC_100 Helicolenus percoides F026 0.559 DC_200 Dendronephthea sp. I111 3.606 DC_100 Hemiasterella sp. 1 I105 14.362 DC_200 Dictyoceratid sp. 1 I100 8.372 DC_100 Herdmania momus I250 6.364 DC_200 Dictyoceratid sp. 10 I163 4.315 DC_100 Ircinia sp. 2 I256 5.848 DC_200 Dictyoceratid sp. 2 I101 9.016 DC_100 Isidid sp. 2 I240 0.344 DC_200 Dictyoceratid sp. 3 I121 3.156 DC_100 Jaspis lutea I099 2.752 DC_200 Dictyoceratid sp. 4 I126 1.288 DC_100 Jaspis stellifera I056 0.258 DC_200 Dictyoceratid sp. 5 I129 0.708 DC_100 Mytilid sp. I254 0.043 DC_200 Dictyoceratid sp. 6 I130 26.597 DC_100 Oceanapia sp. 4 I243 28.208 DC_200 Dictyoceratid sp. 7 I151 0.515 DC_100 Phakellia sp. I116 5.934 DC_200 Dictyoceratid sp. 8 I152 9.660 DC_100 Poecilosclerid sp. I259 10.578 DC_200 Dictyoceratid sp. 9 I154 0.129 DC_100 Polymastia sp. 2 I133 80.711 DC_200 Dictyonella sp. I159 0.708 DC_100 Psammocinia sp. I165 2.322 DC_200 Didemnid sp. 1 I160 0.773 DC_100 Pteronisis sp. 1 I092 0.129 DC_200 Fasciospongia sp. I094 21.703 DC_100 Ptilometra sp. I232 0.645 DC_200 Geodiid sp. I102 87.906 DC_100 Pyxipoma weldii I272 12.169 DC_200 Goniocidaris tubaria I104 13.266 DC_100 Raspailia sp. I125 3.225 DC_200 Hemiasterella sp. 1 I105 32.522 DC_100 Semipallium aktinos I246 0.043 DC_200 Hippospongia sp. I117 5.474 DC_100 Sertularia sp. I091 0.258 DC_200 Hornera robusta I164 0.064 DC_100 Spheciospongia papillosa I109 100.147 DC_200 Ircinia sp. 1 I141 5.796 DC_100 Spheciospongia sp. I097 81.012 DC_200 Isidid sp. 1 I162 0.052 DC_100 Spirophorid sp. I234 17.845 DC_200 Jaspis lutea I099 14.619 DC_100 Stelletta sp. I106 0.172 DC_200 Jaspis stellifera I056 19.320 DC_100 Suberitid sp. I110 6.794 DC_200 Leiosella sp. I098 8.372 DC_100 Triphyllozoon sp. I135 0.086 DC_200 Neotrigonia sp. I290 0.129 DC_200 Adeona spp. I112 8.887 DC_200 Nephtheid sp. I153 0.129 DC_200 Axinella sp. 1 I096 5.410 DC_200 Oceanapia sp. 1 I114 40.314 DC_200 Axinella sp. 2 I115 14.232 DC_200 Oceanapia sp. 2 I128 0.580 DC_200 Bryozoan sp. 1 I131 9.724 DC_200 Oceanapia sp. 3 I150 12.429 DC_200 Bryozoan sp. 2 I146 0.129 DC_200 Ophuiroid sp. 1 I143 0.129 DC_200 Chalinid sp. I095 29.495 DC_200 Ophuiroid sp. 2 I144 0.708 DC_200 Clathria sp. 1 I247 0.451 DC_200 Orthoscuticella spp. I142 1.288 DC_200 Clathrozoon wilsoni I093 4.379 DC_200 Phakellia sp. I116 1.095

SARDI Aquatic Sciences Report – Page 48 Currie, D.R. and Sorokin, S.J. (2011) Canyon biodiversity

Station Species Code Weight Station Species Code Weight DC_200 Polymastia sp. 1 I113 13.717 DC_500 Demosponge sp. 22 I205 0.060 DC_200 Polymastia sp. 2 I133 13.524 DC_500 Galathea sp. I196 0.006 DC_200 Psammocinia sp. I165 1.610 DC_500 Hoplichthys haswelli F028 0.420 DC_200 Pteronisis sp. 1 I092 0.129 DC_500 Hydroid sp. 3 I208 0.006 DC_200 Pyxipoma weldii I272 135.691 DC_500 Ircinia sp. 1 I141 0.040 DC_200 Raspailia sp. I125 4.766 DC_500 Majid sp. I195 0.140 DC_200 Rhizaxinella sp. I127 1.417 DC_500 Nephtheid sp. I153 0.080 DC_200 Sertularia sp. I091 0.193 DC_500 Ophiothrix sp. I198 0.060 DC_200 Siphonochalina sp. I124 0.451 DC_1000 Astropecten sp. 1 I008 0.200 DC_200 Sphaeromid sp. I158 0.064 DC_1000 Bryozoan sp. 4 I173 0.010 DC_200 Spheciospongia papillosa I109 6.698 DC_1000 Bryozoan sp. 8 I214 0.180 DC_200 Spheciospongia sp. I097 67.169 DC_1000 Demosponge sp. 23 I212 0.300 DC_200 Stelletta sp. I106 0.386 DC_1000 Lucinoma galatheae I170 0.060 DC_200 Strongylacidon sp. I132 4.122 DC_1000 Maldanid sp. 2 I176 0.008 DC_200 Suberitid sp. I110 97.115 DC_1000 Serpulid sp. 1 I166 0.010 DC_200 Taonura sp. I122 4.766 DC_1000 Thalacian sp. 1 I213 0.060 DC_200 Triphyllozoon sp. I135 0.258 DC_1500 Actiniarid sp. 1 I172 0.016 DC_200 Umbellulifera sp. I090 49.652 DC_1500 Astropecten sp. 1 I008 0.220 DC_500 Actiniarid sp. 2 I201 0.040 DC_1500 Bryozoan sp. 3 I168 0.004 DC_500 Alcyonacid sp. 1 I206 0.080 DC_1500 Bryozoan sp. 4 I173 0.010 DC_500 Anomiid sp. I200 0.016 DC_1500 Chaetopterid sp. I167 0.008 DC_500 Ascidian sp. 1 I202 0.020 DC_1500 Demosponge sp. 15 I174 0.016 DC_500 Brissus sp. I194 0.420 DC_1500 Demosponge sp. 16 I175 0.018 DC_500 Bryozoan sp. 7 I207 0.018 DC_1500 Hydroid sp. 1 I169 0.012 DC_500 Caryophyllia planilamellata I199 0.720 DC_1500 Lucinoma galatheae I170 0.060 DC_500 Demosponge sp. 14 I161 2.400 DC_1500 Maldanid sp. 1 I177 0.004 DC_500 Demosponge sp. 19 I197 1.160 DC_1500 Maldanid sp. 2 I176 0.008 DC_500 Demosponge sp. 20 I203 0.100 DC_1500 Oculinid sp. I171 0.240 DC_500 Demosponge sp. 21 I204 0.240 DC_1500 Serpulid sp. 1 I166 0.018

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