St Helena Bay Water Quality Trust

ST HELENA BAY WATER QUALITY TRUST:

St Helena Bay State of the Bay 2012

Prepared by

ANCHORenvironmental

St Helena Bay State of the Bay 2012

Prepared for:

St Helena Bay Water Quality Trust Andre du Toit PO Box 655 Veldrif 7635 Tel: 022 7832860 Mobile: 083 2511451 Email: [email protected]

Prepared by:

ANCHOR environmental 8 Steenberg House, Silverwood Close, Tokai 7945, South Africa Tel: 021 701 3420, Fax: 0865428711 www.anchorenvironmental.co.za

Authors: K.L. Tunley, B.M. Clark and A. Biccard

September 2012

Executive summary

Introduction

Regular long-term monitoring in St Helena Bay was initiated by the St Helena Bay Water Quality Trust in 2001. The monitoring programme focuses on water and sediment quality and biotic indices of health, and was designed to provide an overview of trends in the health of the bay, and track to changes that may be caused by human activities. Sediment quality and benthic macrofaunal communities are monitored approximately every five years as part of this programme. The results of these assessments are used to make recommendations regarding the sustainable management of activities in the bay.

This report presents results on sediment characteristics, the levels of organic material in the sediments, and the abundance and distribution of benthic macrofauna living in the sediments of the bay in 2012. It is the third assessment conducted as part of the State of the Bay monitoring program.

Results from the sediment monitoring programme

Sediment and benthic macrofauna samples were collected from 27 sites in St Helena Bay over a 3 day period during the month of April 2012. Samples were collected using a Van Veen grab operated from a Rigid Inflatable Boat (RIB). Samples were analysed for grain size distribution (percentage mud, sand and gravel) and organic content (Total Organic Carbon TOC and Total Organic Nitrogen TON). Assessments of the spatial distribution of different particle size groups and organic content in 2012 were conducted. These were compared with results from 2001 and 2007.

Sediment samples collected in 2012 were comprised predominantly of sand, with a small proportion of mud and no gravel. Relative to data from 2001 and 2007, the 2012 results indicate a dramatic reduction in the amount of mud in the sediments and a significant reduction in the coarser gravel particle fraction in the bay since 2007. This may be associated with an observed increasing intensity and frequency of large storm events over the period 1996 to 2010.

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Data from 2012 also suggest that the spatial distribution of mud in the Bay has also changed dramatically over time, with areas exhibiting the highest mud content having shifted from the northerly and southerly reaches of the Bay in 2007 to the south western edge near the Berg River Estuary mouth in 2012. There have been minor reductions in the levels of organic nitrogen in the sediments at most sites since 2007, while organic carbon levels decreased in the northern reaches, increased in the vicinity of the fish factories and underwent minor changes elsewhere in the bay. The sites in the immediate vicinity of the fish factory outfalls are clearly carbon enriched compared to other areas in the bay. Effluent from fish factories is thus still making an important contribution to organic loading in the sediments but this fortunately is localised to the immediate vicinity (within 100m) of the fish factory outfalls.

Carbon: Nitrogen ratios throughout the bay have increased. These changes may be associated with an increase in denitrification. Denitrification is the breakdown of organic nitrates into nitrogen and ammonia by denitrifying bacteria. These denitrifying bacteria dominate when oxygen availability is low and nitrates are available. This suggests that low oxygen conditions may have prevailed within the benthic environment of the bay since the previous 2007 survey. These conditions may be a result of variations in phytoplankton productivity associated with fluctuations in upwelling-favourable winds. These winds reached a peak in 2001 and underwent a decline up until 2006. Changes in the Southern Oscillation Index (i.e. the oscillation between El Niño and La Niña conditions) may also have played a role.

Sediment conditions in the vicinity of the fish factory outfalls have deteriorated markedly since 2007. These sites are clearly carbon enriched compared to other areas in the bay, and levels of enrichment at these sites have increased disproportionately compared to other sites in the bay. This suggests that the effluent from fish factories is still having a marked effect on the organic composition of the sediments at a local scale (sites immediately adjacent to fish factory outfalls) but not at a large scale.

Results from the benthic macrofauna monitoring programme

All benthic macrofauna (organisms >1 mm in size) in sediment samples collected from the bay were identified to species level, counted, weighed, and assigned to major functional groups (filter feeders, detritivores, predators, scavengers, and grazers). Statistical analyses were performed on these data to assess spatial variability in the benthic macrofauna community structure and composition between sites in 2012, and to assess changes in benthic community structure over time (i.e. in relation to the 2001 and 2007 surveys). Data from 2012 suggest that the benthic macrofaunal community in St Helena Bay was dominated in terms of abundance by the polychaete Diopatra monroi and the amphipod Ampelisca anomala, and in terms of biomass, by the clam Venerupis corrugata and the polychaete D. monroi. Filter feeders were the dominant functional group in terms of biomass in 2012, while filter feeders, detritivores and predators all contributed in relatively similar proportions to total abundance. The remaining groups (scavengers and grazers) contributed relatively little to either biomass or abundance in 2012.

From a spatial perspective, benthic macrofauna communities at sampling stations in the Berg estuary were significantly different from those in the rest of the bay. There were also some

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significant differences among the sampling stations in the bay, with those from the eastern side of the bay being distinct from those in the northern and western sectors, and those taken from the immediate vicinity of the fish factory outfalls being distinct from one another and from other stations in the bay. Abundance of macrofauna in the stations close to the fish factory outfalls was on the whole much lower than those in the rest of the bay with the samples taken in the proximity of the factory in Stompneusbaai being completely devoid of any macrofauna. Given that the macrofauna on the west coast are typically opportunistic species, able to recover rapidly from disturbances and that the communities at two stations were in such a poor state, it is evident that effluent from these particular fish factories are having a negative impact upon the benthic macrofaunal community at these sites. However, as the results of the 2007 survey indicated, these impacts are limited to the local area around the fish factories discharge points.

From a temporal perspective, the overall abundance and biomass of macrofauna in the bay decreased between 2001 and 2007 and then increased between 2007 and 2012. The overall abundance of macrofauna in the bay reached the highest levels on record in 2012, being higher than levels recorded in 2001. Changes between 2007 and 2012 were mostly associated with an increase in the abundance of polychaetes and crustaceans (most notably amphipods) and a change from a community dominated by detritivores to one with a fairly equal proportion of predators, detritivores and filter feeders. Diversity values showed an opposite pattern to that of biomass and abundance with diversity increasing between 2001 and 2007 and then decreasing again by 2012. These fluctuations in the macrofaunal community are likely to be in response to large scale natural disturbances, with few pioneering species dominating the benthos in high numbers following disturbance. Low oxygen events resulting from high levels of productivity are an example of a large scale natural disturbance that is likely to have resulted in these fluctuations in St Helena Bay.

By contrast, macrofauna abundance at the stations in the estuary decreased between 2007 and 2012 and shifted from one dominated by detritivores in 2007 to one dominated by filter feeders in 2012. The cause of this was not clear though and may be related to changes in freshwater runoff to the estuary.

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TABLE OF CONTENTS

Executive summary ...... i TABLE OF CONTENTS ...... iv 1 INTRODUCTION ...... 1 2 STATE OF THE BAY MONITORING ...... 2 3 SAMPLING METHODS...... 3 4 SEDIMENT ...... 6 4.1 Particle size ...... 7 4.2 Organic matter ...... 10 4.2.1 Total Organic Carbon ...... 10 4.2.2 Total Organic Nitrogen ...... 12 4.2.3 Carbon to nitrogen ratios ...... 14 5 BENTHIC MACROFAUNA ...... 16 5.1 Methods ...... 16 5.1.1 Laboratory Analysis ...... 16 5.1.2 Statistical Analysis ...... 16 5.2 Results ...... 18 5.2.1 Community structure and composition ...... 18 5.2.2 Diversity indices ...... 31 6 INTEGRATION OF PHYSICO-CHEMICAL PARAMETERS AND BIOTIC INDICATORS ...... 33 6.1 Methods ...... 33 6.2 Results ...... 33 7 DISCUSSION ...... 35 7.1 Bay-wide variability ...... 35 7.2 Fish factory sites ...... 38 7.3 Estuarine sites ...... 38 8 CONCLUSIONS ...... 40 9 REFERENCES ...... 41

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1 INTRODUCTION

St Helena Bay is situated on the west coast of South Africa and extends from Dwarskersbos in the north, past the Shelley Point peninsula, to Cape St Martin in the west, encompassing 18 smaller bays and the estuary of the Berg River. The Bay is positioned in the southern section of the Benguela Current System (BCS) which extends along the west coast of southern Africa between Cape Agulhas (South Africa) and the Congo River mouth (Angola). The BCS is one of four major eastern-boundary current systems which is characterised by the wind-driven upwelling of cold, nutrient rich water (Shannon & O’Toole 1998). This cold current originates from the South Atlantic Circulation, which circles just north of the Arctic Circumpolar Current. The naturally cool temperature of the Benguela current (average temperature 10-14oC) is enhanced by the upwelling of the even cooler nutrient-rich deep water (Branch 1981). The area experiences strong southerly and south-easterly winds which are deflected by the Coriolis forces (rotational force of the earth which causes objects in the southern hemisphere to spin anticlockwise). These prevailing conditions deflect the surface waters offshore and cause cold, nutrient rich water to upwell against the shore . This water is the nutrient rich life force of the west coast. Phytoplankton bloom when the nutrients reach the surface waters where plenty of light is available for photosynthesis, and the phytoplankton is then preyed upon by zooplankton, which is in turn eaten by filer feeding fish such as anchovy or sardine. This makes the west coast one of the richest fishing grounds in the world and also attracts large colonies of sea birds and seals (Branch 1981).

St Helena Bay is positioned downstream of the Cape Columbine upwelling cell and is a retention zone for the nutrient rich water that is upwelled in this cell (Monteiro and Roychoudhury 2005). St Helena Bay functions as a productive nursery to early life stages of fish (Bakun 1998, Monteiro and Roychoudhury 2005), however it is also subject to incidence of harmful algal blooms and regular episodes of oxygen depletion in the coastal waters, which have in the past lead to major mortality events for organisms such as rock lobsters and fish (Cockroft et al. 2000, Monteiro & Roychoudhury 2005). A rock lobster reserve has also been declared within the bay in terms of the Marine Living Resources Act. The Berg River Estuary, which is considered part of the Bay, is one of a few estuaries draining off the west coast and is ecologically important for both fisheries and birds. Tourism is also becoming an increasingly important industry supported by the bay given its picturesque and sheltered nature and the variety of recreational opportunities it offers including sailing, canoeing, surfing, bathing, diving, kitesurfing, fishing and beach activities.

St Helena Bay was the centre of the pelagic fishing industry when it began in the 1940s and has been an important part of the West Coast lobster fishing grounds (von Bonde & Merchand 1935, Hutchings et. al. 2012). The Bay still functions as a major node for industrial and small-scale fisheries on the west coast. Indeed, the fishing industry is the mainstay of St Helena Bay with the bulk of South Africa’s fish production and processing being conducted in the St Helena Bay factories. The factories in the bay produce a variety of products including tinned fish, fresh and frozen hake, fish- meal and live crayfish for export. Other industries located around the periphery of the bay include mariculture, ship repair and shipbuilding.

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2 STATE OF THE BAY MONITORING

Regular, long-term environmental monitoring is essential to identify impacts and to implement measures to minimise negative human impacts on the environment (e.g. pollution) and maintain the beneficial value of an area. This is particularly important to an area such as St Helena Bay, which serves as a major node for industrial as well as small scale fisheries, while at the same time supporting important ecological functions and a growing tourism industry. Fish processing and vessel operations in St Helena Bay have in the past been shown to impact marine biota and water quality in the Bay, while other industrial, residential and tourism developments also have the potential to negatively impact on ecosystem health of the Bay. Various methods are available to monitor the health of the environment, including the measurement of physical parameters (e.g. water temperature, oxygen levels, and circulation patterns), actual pollutants in the water column, sediments or tissue of biota (e.g. heavy metals, hydrocarbons, microbiological indicators) and biological components of the ecosystem (e.g. birds, fish, and invertebrates).

Organic matter is one of the most universal pollutants affecting marine life and if it is allowed to accumulate to high levels or if it is introduced into the environment at a rate faster than it can be assimilated, it can lead to significant community disturbance. Organic loading is a particular concern for St Helena Bay given the number of fish processing facilities operational in the bay and the fact that the bay is a retention zone. High organic loading typically leads to eutrophication, which may bring about a number of community responses. These include increased growth rates by primary producers, disappearance of organisms due to hypoxia or anoxia, changes in community composition and reduction in the number of species following repeat hypoxia and even complete disappearance of benthic organisms in severely eutrophic and anoxic sediments (Warwick 1993).

It is important to monitor biological criteria in addition to physico-chemical and ecotoxicological variables as biological indicators provide a direct measure of the state of the ecosystem. Benthic macrofauna are the biotic component most frequently monitored to detect changes in the health of the marine environment. This is primarily because these species are short lived and therefore their community composition responds noticeably to changes in environment quality over time (Warwick 1993). Given that they are also relatively non-mobile (as compared with fish and birds) they tend to be directly affected by pollution and they are easy to sample quantitatively (Warwick 1993). Furthermore they are well-studied scientifically, in comparison to other sediment-dwelling components (e.g. meiofauna and microfauna) and taxonomic keys are available for most groups. In addition, community response to a number of anthropogenic influences has been well documented.

The major industries as well as other stakeholder in the St Helena Bay area are represented on the St Helena Bay Water Quality Trust. The Trust has established a long term monitoring programme, which monitors water quality, sediment quality and biotic indices of health, to determine the overall health of the bay and track changes that may be caused by human activities. Sediment quality and benthic macrofaunal communities are monitored approximately every five years. The results of these assessments are used to make recommendations regarding the sustainable management of activities in the bay.

This report presents results on sediment characteristics, the levels of organic material in the sediments, and the abundance and distribution of benthic macrofauna living in the sediments of the

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bay in 2012. It is the third assessment conducted as part of the State of the Bay monitoring program which commenced in 2001. The methods used in this assessment replicated the sampling approach used in previous State of the Bay surveys as closely as possible, to minimise any variations in macrofauna abundance or composition that may be introduced from this source.

3 SAMPLING METHODS

Previously (2001 and 2007) sediment samples were collected from 33 sites across several months period (April-August) throughout the Bay using a van Veen Grab or an Ocean Instruments multicorer. All samples were analysed for trace metal content, acid volatile sulphides, sediment particle size and particulate organic carbon and nitrogen. Benthic macrofauna samples were collected at 17 sites distributed throughout the Bay using a Van Veen grab operated from a Rigid Inflatable Boat (RIB). The dimensions of the grab used were 0.33 x 0.33 m resulting in a sampled area of ~0.1 m2/grab. The sediment was washed through a 1 mm mesh sieve. Benthic macrofauna retained on the sieve was washed into labelled sample bottles, fixed in ~4% formaldehyde solution, and transferred to the laboratory for processing. One replicate was taken at the majority of sites in the Bay and multiple (3 – 5) replicates were taken at sites in the vicinity of the fish farms. Sites were sampled over three days spread between April and July of 2007.

In this study (2012) benthic macrofauna and sediment samples were collected from 27 sites throughout St Helena Bay over a 3 day period during the month of April. Sampling was conducted using a Van Veen grab operated from a RIB. The dimensions of the grab used were 0.34 x 0.42 m resulting in a sampled area of ~0.143 m2/grab. A subsample from each grab was collected and used for sediment texture and organic content analysis and the balance of the sediment was washed through a 1 mm mesh sieve. Benthic macrofauna retained on the sieve were washed into labelled sample bottles, fixed in ~4% formaldehyde solution, and transferred to the laboratory for further processing.

Figure 1: The van Veen Grab used for sediment and benthic macrofauna sample collection in 2012 and the filtering of sediments through a 1mm mesh bag.

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Figure 2. Sites sampled in 2012 for sediment and benthic macrofauna as part of the State of the Bay survey. The position of sites relative to fish processing facilities outlets (black squares) is indicated.

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Table 1 Sites sampled for benthic macrofauna and sediment in St Helena Bay during the 2001, 2007 and 2012 State of the Bay surveys.

2001 2007 2012 Site Macrofauna and Lat (S) Lon (E) Name Macrofauna Sediment Macrofauna Sediment Sediment 32.6950 17.9250 SH3 X X Not sampled X Not sampled 32.6878 17.9255 SH4 X X Not sampled X X 32.5502 17.9243 SH6 X X X X X 32.6727 18.0237 SH11 X X Not sampled X Not sampled 32.7078 18.0240 SH12 X X X X X 32.7252 18.0233 SH13 X X Not sampled X X 32.7498 18.0280 SH14 X X Not sampled X X 32.7592 18.0403 SH15 X X Not sampled X X 32.7632 18.0492 SH16 X X Not sampled X X 32.7250 18.1238 SH21 X X X X X 32.4245 18.1248 SH26 X X X X X 32.4248 18.2492 SH27 X X Not sampled X X 32.6007 18.2485 SH29 X X X X X 32.6330 18.2480 SH30 X X X X X 32.6587 18.2355 SH31 X X X X X 32.7673 18.0583 SH33 X X Not sampled X X 32.7510 18.0582 SH34 X X Not sampled X Not sampled 32.7257 18.0590 SH35 X X Not sampled X Not sampled 32.7230 17.9888 SH36 X X Not sampled X X 32.7000 17.9500 SH40 Not sampled Not sampled Not sampled X Not sampled 32.7439 18.0191 SH45 Not sampled Not sampled Not sampled X X 32.7766 18.1363 SH46 Not sampled Not sampled X X X 32.7806 18.1460 SH47 Not sampled Not sampled X X X 32.7907 18.1465 SH48 Not sampled Not sampled X X X 32.7881 18.1687 SH49 Not sampled Not sampled X X X 32.6968 18.1960 SH50 X (40) X (40) X X X 32.7180 17.9437 SH51 X (37) X (37) Not sampled X X 32.7169 17.9342 SH52 Not sampled Not sampled Not sampled X Not sampled 32.7457 18.0151 SH53 Not sampled Not sampled X (SH166) X X 32.7478 18.0145 SH54 Not sampled Not sampled X (SH167) X X 32.7736 18.0514 SH55 Not sampled Not sampled X (SH168) X X 32.7753 18.0513 SH56 Not sampled Not sampled X (SH168) X X 32.7233 17.9778 SH57 Not sampled Not sampled X (SH170) X X

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4 SEDIMENT

The sediment samples were tested for Total Organic Carbon (TOC) and Total Organic Nitrogen (TON) by the CSIR using the standard laboratory method CSIR (MALS 3.3) (CSIR 2008). The particle size composition was determined by Scientific Services using the following sieve mesh sizes: 2000 µm, 1000 µm, 850µm, 710µm, 500µm, 425µm, 300µm, 250µm, 125µm and 63µm.

The assessment of these results considered both the spatial distribution of different particle size groups and organic content as well as the changes in distribution and the extent of organic loading in 2012 relative to 2001 and 2007. TOC and TON were however not assessed in 2001 and therefore the temporal assessment of organic loading was only conducted for 2007 and 2012.

Table 2. Sediment particle size composition and organic content results for St Helena Bay in 2012. C:N ratios presented in this table are based on molar mass. TOC TON Site Depth (m) Gravel (%) Sand (%) Mud (%) (ppm) (ppm) C:N SH 4 31.0 0.00 92.45 7.55 2.98 0.30 11.61 SH 6 66.0 0.00 99.84 0.16 0.27 0.02 19.61 SH 12 20.0 0.00 90.91 9.09 4.36 0.46 11.14 SH 13 15.0 0.00 89.75 10.25 3.74 0.41 10.68 SH 14 8.0 0.00 95.48 4.52 3.61 0.36 11.80 SH 15 6.0 0.00 96.14 3.86 3.89 0.40 11.30 SH 16 5.0 0.00 82.84 17.16 2.59 0.27 11.12 SH 21 8.0 0.00 94.33 5.67 0.60 0.04 17.41 SH 26 72.0 0.00 97.59 2.41 4.50 0.44 11.95 SH 27 48.0 0.00 98.94 1.06 1.07 0.08 15.40 SH 29 17.0 0.00 98.41 1.59 0.17 0.01 39.67 SH 30 10.0 0.00 99.62 0.38 0.32 0.02 21.00 SH 31 6.5 0.00 99.19 0.81 0.23 0.01 24.29 SH 33 5.0 0.00 67.95 32.05 3.24 0.32 11.79 SH 36 8.0 0.00 93.06 6.94 1.86 0.18 12.16 SH 46 1.0 0.00 83.93 16.07 0.90 0.07 14.42 SH 47 1.0 0.00 99.35 0.65 0.31 0.01 29.94 SH 48 1.0 0.00 96.47 3.53 0.54 0.04 18.00 SH 49 1.0 0.00 97.32 2.68 0.43 0.03 20.25 SH 50 5.0 0.00 84.77 15.23 0.45 0.03 16.26 SH 51 20.0 0.00 99.30 0.70 8.54 0.43 23.23 SH 53 1.0 0.00 98.54 1.46 7.16 0.71 11.75 SH 54 1.0 0.00 94.20 5.80 7.11 0.60 13.91 SH 55 4.0 0.00 83.57 16.43 2.56 0.15 19.63 SH 56 6.0 0.00 99.88 0.12 3.59 0.08 52.34 SH 57 2.0 0.00 96.51 3.49 3.78 0.35 12.46

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4.1 Particle size

The sediments sampled at all sites in St Helena Bay in 2012 were comprised predominantly of sand, with a small proportion of mud and no gravel. The sites situated in the Bay in close proximity to the Berg River Estuary mouth had the highest proportion of mud (Figure 3 and Figure 6). Relative to data from 2001 and 2007, the 2012 results indicate a dramatic reduction in the amount of mud in the sediments in St Helena Bay (Figure 3, Figure 4 and Figure 5). In addition, there has been a significant reduction in the coarser gravel particle fraction in the bay since 2007 (Figure 4), making the situation in 2012 (Figure 3) more similar to that evident in 2001 (Figure 5). The spatial distribution of mud in the Bay has also changed dramatically over time, with areas exhibiting the highest mud content having shifted from the northerly and southerly reaches of the Bay in 2007 (Figure 7) to the south western edge near the Berg River Estuary mouth (Figure 6).

100% 90% 80% 70% 60% 50% Mud 40% Sand 30% Gravel 20% 10%

SH 4 4 SH SH 6 6 SH

SH 12 12 SH 14 SH SH 13 13 SH 15 SH 16 SH 21 SH 26 SH 27 SH 29 SH 30 SH 31 SH 33 SH 36 SH 46 SH 47 SH 48 SH 49 SH 50 SH 51 SH 53 SH 54 SH 55 SH 56 SH 57 SH

Figure 3: Particle size composition of sediment sampled at sites in 2012.

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100% 90% 80% 70% 60% 50% Mud 40% Sand 30% Gravel 20% 10%

SH 4 4 SH 6 SH

SH 46 46 SH 48 SH 50 SH 53 SH SH 13 13 SH 14 SH 15 SH 16 SH 21 SH 26 SH 27 SH 29 SH 30 SH 31 SH 33 SH 36 SH 47 SH 49 SH 51 SH 54 SH 55 SH 56 SH 57 SH SH 12 12 SH

Figure 4. Particle size composition of sediment sampled at sites in 2007.

100% 90% 80% 70% 60% 50% Mud 40% Sand 30% Gravel 20% 10%

SH 4 SH SH 6 SH

SH 13 SH 15 SH 16 SH 26 SH 27 SH 30 SH 36 SH 47 SH 48 SH 50 SH 51 SH 54 SH 57 SH SH 14 SH 21 SH 29 SH 31 SH 33 SH 46 SH 49 SH 53 SH 55 SH 56 SH SH 12 SH

Figure 5. Particle size composition of sediment sampled at sites in 2001.

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Figure 6. Spatial distribution of mud interpolated from sites sampled in St Helena Bay in 2012.

Figure 7. Spatial distribution of mud interpolated from sites sampled in St Helena Bay in 2007.

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4.2 Organic matter

4.2.1 Total Organic Carbon

The results of the TOC analysis for 2012 revealed that the area in the vicinity of the fish factories and in Britannia Bay (southern reaches of the bay) is highly enriched in carbon in comparison to other areas in the Bay. One site on the northern edge of the Bay (SH26, the deepest site sampled) also indicated a moderate to high level of carbon enrichment (Figure 8 and Figure 10 ). The spatial distribution of TOC through the bay did not correlate with that of mud.

A temporal comparison between the TOC results of 2007 and 2012 suggests that there had been a reduction in the levels of carbon enrichment in the northern reaches of the Bay (SH26) and an increase in the vicinity of the fish factories (SH55, SH56 and SH57) and in Britannia Bay (SH51) (Figure 8, Figure 9 and Figure 10). Minor changes in carbon enrichment levels (predominantly slight reductions) were evident at all other sites in the Bay and in the estuary.

Figure 8. Spatial distribution of Total Organic Carbon (TOC) interpolated from sites sampled in St Helena Bay in 2012.

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Figure 9: Spatial distribution of Total Organic Carbon (TOC) interpolated from sites sampled in St Helena Bay in 2007.

9.00 8.00 7.00

6.00 5.00 4.00 2007

TOC (ppm) 3.00 2012 2.00 1.00

0.00

SH 4 4 SH 6 SH

SH 13 13 SH 47 SH SH 14 14 SH 15 SH 16 SH 21 SH 26 SH 27 SH 29 SH 30 SH 31 SH 33 SH 36 SH 46 SH 48 SH 49 SH 50 SH 51 SH 53 SH 54 SH 55 SH 56 SH 57 SH SH 12 12 SH

Figure 10. Total Organic Carbon in sediments in St Helena Bay in 2007 and 2012.

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4.2.2 Total Organic Nitrogen

The results of the TON analysis for 2012 revealed that the area in the vicinity of the St Helena Bay harbour and the fish factory outlets (southern reaches of the bay) is highly enriched in nitrogen in comparison to other areas in the Bay. The deepest station on the northern edge of the Bay (SH26) also exhibited a moderate to high level of nitrogen enrichment.

A temporal comparison between the TON results of 2007 and 2012 revealed that there had been a reduction in the levels of nitrogen enrichment at all sites in the Bay with the exception of the sites in the vicinity of the fish factory in Stompneusbaai (SH57) and in Britannia Bay (SH 51) where the nitrogen content had increased since 2007.

1.40 1.20

1.00 0.80 2007 0.60

TON(ppm) 2012 0.40 0.20

0.00

SH 4 4 SH 6 SH

SH 12 12 SH 46 SH SH 14 14 SH 15 SH 16 SH 21 SH 26 SH 27 SH 29 SH 30 SH 31 SH 33 SH 36 SH 47 SH 48 SH 49 SH 50 SH 51 SH 53 SH 54 SH 55 SH 56 SH 57 SH SH 13 13 SH

Figure 11. Total Organic Nitrogen in sediments in St Helena Bay in 2007 and 2012.

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Figure 12. Spatial distribution of total organic nitrogen (TON) interpolated from sites sampled in St Helena Bay in 2012.

Figure 13. Spatial distribution of total organic nitrogen (TON) interpolated from sites sampled in St Helena Bay in 2007.

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4.2.3 Carbon to nitrogen ratios

All the sites sampled in 2012 had high C:N ratios. Three sites where the ratios were exceptionally high included SH56 (adjacent to a fish factory), SH29 (eastern inshore area) and SH47 in the estuary (Table 3). However of these three sites, only SH 56 was highly enriched with carbon, while the other two sites had relatively low carbon concentrations (Figure 8). A comparison between the 2007 and 2012 results revealed an increase in the C:N ratios at all sites in the Bay since 2007.

Table 3: C:N ratios at all sites sampled in St Helena Bay in 2007 and 2012.

Area Site 2007 2012 Britannia Bay SH 51 7.9 23.2 SH 53 9.2 11.7 SH 54 7.8 13.9 Fish Factories SH 55 7.3 19.6

SH 56 6.5 52.3 SH 57 7.0 12.5 Western reaches SH 4 7.4 11.6 South Western reaches SH 6 5.0 19.6 SH 12 8.2 11.1 SH 13 7.9 10.7 SH 14 7.9 11.8 SH 15 8.1 11.3 Southern reaches SH 16 7.7 11.1 SH 21 6.7 17.4 SH 33 7.8 11.8 SH 36 7.9 12.2 SH 29 6.6 39.7 SH 30 6.0 21.0 Eastern inshore SH 31 4.7 24.3 SH 50 5.3 16.3 SH 26 9.4 11.9 Northern Reaches SH 27 9.3 15.4 SH 46 6.9 14.4 SH 47 6.6 29.9 Berg River Estuary SH 48 7.2 18.0 SH 49 7.0 20.3

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Figure 14. C:N ratio interpolated from sites sampled in St Helena Bay in 2012.

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5 BENTHIC MACROFAUNA

5.1 Methods

5.1.1 Laboratory Analysis

In the laboratory, benthic macrofauna samples were washed to remove all traces of formaldehyde, and transferred to 1% phenoxatol (ethylenglycolmonophenyl-ether). Thereafter all animals considered alive at the time of collection (i.e. excluding empty shells) were identified to the lowest taxon possible, blot dried and weighed to the nearest 0.001 g on a precision balance. The reference collection from the 2007 survey was sourced and compared to that of the 2012 survey to ensure consistency in the species identification. A majority of the species collected in 2001 were not identified to a species level and no reference collection was available for this survey.

5.1.2 Statistical Analysis

The principle aim of monitoring the health of an area is to detect the effects of stress, as well as to monitor recovery after an environmental perturbation. There are numerous indices, based on benthic invertebrate fauna information, which can be used to reveal conditions and trends in the state of ecosystems. These indices include those based on community composition, diversity and species abundance and biomass. Given the complexity inherent in environmental assessment it is recommended that several indices be used (Salas et al. 2006). The community composition, diversity, and species abundance and biomass of soft bottom benthic macrofauna samples, collected in St Helena Bay in 2012, are considered in this report.

The data collected from this survey were used for two purposes

1) to assess spatial variability in the benthic macrofauna community structure and composition between sites in 2012 and

2) to assess changes in benthic community structure over time (temporal variation) (i.e. in relation to the 2001 and 2007 surveys).

Both the spatial and temporal assessments are necessary to provide a good indication of the state of the system.

Community structure and composition The statistical program PRIMER 6 (Clarke and Warwick 1993) was used to conduct a spatial assessment of the benthic macrofauna data collected in 2012 and a temporal comparison between the macro benthic communities sampled in 2007 and 2012. The temporal comparison did not include the 2001 data set as no reference collection was sourced and consistency in the identification of species could not be guaranteed. Data were root-root (fourth root) transformed and converted to a similarity matrix using the Bray-Curtis similarity coefficient. A cluster analysis was performed in order to find ‘natural groupings’ between samples (sites). The results of the cluster analysis are displayed on a dendrogram which graphically depicts the similarity among sites

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by clustering them in groups. Statistically significant clusters of sites are revealed using a SIMPROF analysis. These results were plotted geographically using GIS software to reveal any spatial trends in the sites grouped in accordance with community composition similarity. SIMPER analysis was used to identify species principally responsible for the clustering of sites. These results were used to characterise different regions of sites based on the communities present at the sites. It is important to remember that the community composition is a reflection of not only the physico-chemical health of the environment but also the ability of communities to recover from disturbance.

A total of eight sites were sampled across all three surveys - 2001, 2007 and 2012. These sites were all within the bay. No estuary or fish factory sites were sampled in 2001. The abundance and relative proportions of benthic macrofauna taxonomic groups sampled at these eight sites were compared.

Diversity Indices A number of indices (single numbers) can be used as measures of community structure; these include the total number of individuals (N), total number of species (S), the total biomass (B), and the species equability or evenness, which is a measure of how evenly individuals are distributed among different species. Diversity indices provide a measure of diversity, i.e. the way in which the total number of individuals is divided up among different species. Understanding changes in benthic diversity is important because increasing levels of environmental stress generally result in a decrease in diversity.

Two different aspects of community structure contribute to community diversity, namely species richness and equability (evenness). Species richness refers to the total number of species present while equability or evenness expresses how evenly the individuals are distributed among different species. A sample with greater evenness is considered to be more diverse. It is important to note when interpreting diversity values that predation, competition and disturbance all play a role in shaping a community. For this reason it is important to consider physical parameters as well as other biotic indices when drawing a conclusion from a diversity index.

The following measures of diversity were calculated for each sampling location using PRIMER V 6:

The Shannon-Weiner diversity index (H’): H’ = - Σipi(log pi) (1)

where pi is the proportion of the total count arising from the ith species. This is the most commonly used diversity measure and it incorporates both species richness and equability.

The Pielou’s evenness index (J’): J’ = H’observed / H’max (2)

where H’max is the maximum possible diversity which would be achieved if all species were equally abundant (= log S). This is the most common expression of equability.

The Margalef’s index (d) of species richness: D = (S-1)/ log N (3)

where S is the total number of species and N is the total number of individuals.

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Species richness is often simply referred to as the total number of species (S), but this is dependent on sample size. The Margalef’s index thus incorporates the total number of individuals (N) and is a measure of the total number of species present for a given number of individuals.

The diversity (H’) value for each site was plotted geographically and this was used to interpolate values for the entire system in order to reveal any spatial patterns. Eight sites were sampled in all three years. The diversity and abundance values for each of these years was compared.

5.2 Results

5.2.1 Community structure and composition

The benthic macrofaunal community in St Helena Bay in 2012 was dominated in terms of abundance by the polychaete Diopatra monroi and the amphipod Ampelisca anomala and in terms of biomass by the clam Venerupis corrugata (Figure 15, Table 4). The polychaete D. monroi also comprised a large proportion of the total biomass.

100% SIPUNCULIDEA

90% PYCNOGONIDA

80% POLYCHAETA

70% MALACOSTRACA (ISOPODA)

60% MALACOSTRACA (BRACHYURA) MALACOSTRACA (AMPHIPODA) 50% MALACOSTRACA 40% GASTROPODA 30% BRACHIOPODA 20% BIVALVIA 10% ASCIDIACEA 0% Abundance Biomass ANTHOZOA

Figure 15. Relative contributions of taxa to the overall biomass and abundance of benthic macrofauna sampled in St Helena Bay in 2012.

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Table 4: Benthic macrofaunal species comprising >1% of the total count or biomass in the 2012 St Helena Bay survey.

Taxonomic Level Phylum Class Species % Abundance % Biomass Cnidaria Anthozoa Anemone sp. A 2.88 2.16 Anemone sp. B 0.30 1.18 Virgularia schultzei 0.44 6.61 Arthropoda Malacostraca Upogebia africana 0.98 2.04 Ampelisca anomala 23.48 0.31 Corophiid A 6.33 0.07 Ampelisca palmata 3.95 0.06 Melita subchelata 2.40 0.04 Lemboides crenatipalma 1.94 0.03 Ampelisca spinimana 2.46 0.03 Photis longimanus 1.50 0.00 Photis longidactylus 1.44 0.00 Gammaropsis sophiae 1.06 0.00 Mollusca Bivalvia Venerupis corrugata 2.37 35.63 Dosinia lupinus orbignyi 0.90 6.41 Choromytilus meridionalis 0.04 5.02 Tellina gilchristi 1.79 4.52 Macoma crawfordi ordinaria 0.98 2.89 Gastropoda Nassarius vinctus 0.90 1.08 Annelida Polychaeta Diopatra monroi 24.61 18.32 Lumbrinereis heteropoda difficilis 0.50 2.88 Arenicola loveni 0.29 2.59 Pherusa swakopiana 0.62 1.30 Nephtys hombergii 1.02 0.82 Pectinaria capensis 1.16 0.81 Sabellides luderitzi 2.23 0.16 Magelona papillicornis 6.20 0.14

Dominant taxonomic groups included polychaetes and crustaceans (where abundance was considered) and bivalves, polychaetes and Anthozoa (anemones and sea pens) (where biomass was considered)(Figure 15). Filter feeders were the dominant functional group in terms of biomass while filter feeders, detritivores and predators all contributed in relatively similar proportions to the total abundance of the benthic macrofauna in the bay (Figure 16).

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100%

90%

80% SCAVENGER 70% PREDATOR 60%

50% GRAZER

40% FILTER FEEDER

30% DETRITIVORE

20%

10%

0% Abundance Biomass

Figure 16. Relative contributions of functional groups to the overall biomass and abundance of benthic macrofauna sampled in St Helena Bay in 2012.

Spatial variation The benthic macrofaunal communities sampled in the Berg River Estuary (SH46-49) were significantly different to those sampled in the bay (Figure 17). The benthic macrofaunal communities at all four sites in the estuary (Group A on Figure 17) were similar to one another and clustered together in the dendrogram. These estuarine sites were characterised by the presence of the estuarine mud prawn (Upogebia africana), two detritivorous polychaetes species (Orbinia angrapequensis and Nereis spp.) and the three-legged crab (Spiroplax spiralis).

Four of the sites sampled on the inshore eastern edge of the bay were similar to one another (Group B) and were characterised by polychaetes (Magelona papillicornis, Nephtys hombergii, Pectinaria capensis and Sabellides luderitzi), an anemone and the isopod (Anthelura remipes)(Figure 20). These sites were all similar to one of the sites adjacent to a fish factory effluent pipe.

The dendrogram and SIMPROF analysis revealed that three of the sites sampled in the southern reaches of the Bay were similar to one another (Group E) (Figure 20). These sites were characterised by a predatory polychaete (Diopatra monroi), three amphipod species (Ampelisca anomala, Lemboides crenatipalma and a Corophiid spp.) and an anemone species.

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C B D

Figure 17. Dendrogram representing the similarity of sites (Bray Curtis Similarity) based on the benthic macrofaunal community composition sampled in St Helena Bay in 2012. The red lines represent significant clusters based on the SIMPROF analysis

There was no other clear spatial clustering of the remainder of sites in the bay. Sites sampled in the northern reaches of the bay were similar to two sites sampled in the southern reaches of the Bay (Group D) and were characterised by the presence of two polychaetes (Diopatra monroi and Lumbrinereis heteropoda difficulties), the amphipod Ampelisca spinimana and the clam Dosinia lupinus orbignyi (Figure 20). These sites all had a relatively low to moderate abundance and biomass.

All of the sites adjacent to the fish factory effluent pipes were significantly different to one another (Figure 20). One site sampled adjacent to a fish factory was significantly similar to a site sampled in Britannia Bay. No species were recorded in the sample collected at site SH57 adjacent to the fish factory in Stompneusbaai and as a result this site had no similarity to other sites sampled and was excluded from the spatial analysis conducted. Results of the cluster analysis indicated that site SH56, situated adjacent to the most southerly fish factory was another clear outlier from other sites sampled in the Bay, however, three species were recorded (the average number of species per site was 14.46) in very low abundance. The species recorded at this site were all polychaetes with low biomass.

The sites sampled in the southern reaches of the Bay generally had the highest abundance and biomass of benthic macrofauna and those in the estuary generally had the lowest (Figure 18 and Figure 19). There was much variability in the total abundance and biomass of benthic macrofauna between the sites sampled at the fish factories.

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14000

12000

10000

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6000

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Total number Total individuals of per m2 2000

SH 4 SH 6 SH

SH45

SH 57 SH

SH 16 SH 21 SH 33 SH 36 SH 29 SH 30 SH 31 SH 50 SH 26 SH 27 SH 46 SH 47 SH 48 SH 49 SH SH 51 SH 53 SH 54 SH 55 SH 56 SH 12 SH 13 SH 14 SH 15 SH

Brit Fish Factories W S E N Estuary

Figure 18: Total abundance (no. organisms per m2) of benthic macrofauna at sampling stations in St Helena Bay in 2012.

8000 7000

2 6000 5000 4000 3000

2000 Total biomass biomass Total per m 1000

SH 4 SH 6 SH

SH45

SH 57 SH

SH 51 SH 54 SH 55 SH 13 SH 14 SH 21 SH 33 SH 29 SH 50 SH 26 SH 47 SH SH 53 SH 56 SH 12 SH 15 SH 16 SH 36 SH 30 SH 31 SH 27 SH 46 SH 48 SH 49 SH Brit Fish Factories W S E N Estuary

Figure 19. Total biomass (g per m2) of benthic macrofauna at sampling stations in St Helena Bay in 2012.

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Figure 20: Geographic representation of the results of a PRIMER analysis showing similarity of benthic macrofaunal community composition in St Helena Bay in 2012. Sites with similar species are placed in groups demoted by the different colour dots on the map.

POLYCHAETA

Nephtys hombergii (Predator) Glycera convoluta (Predator) Photograph by: N. Steffani Photograph by: N. Steffani

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MALACOSTRACA (Amphipoda)

Hippomedon normalis (Scavenger) Ampelisca brevicornis (Filter feeder) Photograph by: N. Steffani Photograph by: N. Steffani

MALACOSTRACA (Decapoda)

Callichirus kraussi (Detritivore) Upogebia africana (Filter feeder) Photograph by: C. Griffiths Photograph by: C. Griffiths

GASTROPODA

Nassarius vinctus (Scavenger) Venerupis corrugata (Filter feeder) Photograph by: N. Steffani

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Temporal variation (2001-2012: Inner bay only) The abundance of macrofauna at the eight sites sampled in 2001, 2007 and 2012 decreased between 2001 and 2007 and increased again in 2012 (Figure 21).

Polychaetes and malacostraca (most notably the amphipods) dominated at most of these eight sites in all three sampling years. Furthermore, fluctuations in the abundance of polychaetes and malacostraca accounted for most of the changes in overall abundance of benthic macrofauna at the these sites. With the exception of Sites SH31 and SH50, the benthic macrofaunal communities sampled at these eight sites in 2012 resemble that sampled in 2001 in terms of abundance and taxonomic dominance.

Bivalves, anthozoa (anemones) and polychaetes dominated the benthic macrofaunal communities in terms of biomass in 2001 (Figure 22). There was a dramatic reduction in the total biomass sampled at seven of the eight sites between 2001 and 2007 (Site SH29 was the exception). Much of this reduction was accounted for by bivalves and polychaetes. The biomass then increased between 2007 and 2012 at all sites. Much of this reduction was accounted for by bivalves, anthozoa and polychaetes. The benthic macrofauna sampled at these eight sites in 2012 resemble that sampled in 2001 in terms of biomass and taxonomic dominance.

Page | 25 120000 119000 118000

120000117000 119000116000 118000115000 117000114000 116000113000 115000112000

120000114000111000 119000113000110000 118000112000109000 117000111000108000 116000110000107000 115000109000106000 114000108000105000 113000107000104000 112000106000103000 105000111000102000 104000110000101000 103000109000100000 10200010800099000 10100010700098000 10000010600097000 1050009900096000 1040009800095000 1030009700094000 1020009600093000 1010009500092000 1000009400091000 930009900090000 920009800089000 910009700088000 900009600087000 890009500086000 880009400085000 870009300084000 860009200083000 850009100082000 840009000081000 830008900080000 820008800079000 810008700078000 800008600077000 790008500076000 780008400075000 770008300074000 760008200073000 750008100072000 740007100080000 730007000079000 720006900078000 710006800077000 700006700076000 690006600075000 680006500074000 670006400073000 660006300072000 650006200071000 640006100070000 630006000069000 620005900068000 610005800067000 600005700066000 590005600065000

580005500064000 2 570005400063000 560005300062000 550005200061000 540005100060000 530005000059000 520004900058000

510004800057000 Total number of individuals numberTotal individuals of per m 500004700056000 490004600055000 480004500054000 470004400053000 460004300052000 450004200051000 440004100050000 430004000049000 420003900048000 410003800047000 400003700046000 390003600045000 380003500044000 370003400043000 360003300042000 350003200041000 340003100040000 330003000039000 320002900038000 310002800037000 300002700036000 290002600035000 280002500034000 270002400033000 260002300032000 250002200031000 240002100030000 230002000029000 220001900028000 210001800027000 200001700026000 190001600025000

8000 180001500024000 2 7000 170001400023000 12000022000 6000 1600013000 POLYCHAETA PENNATULACEA 11900021000 5000 1500012000 MALACOSTRACA HOLOTHUROIDEA 11800020000 4000 1400011000 GASTROPODA BIVALVIA 19000 3000 1170001300010000 ACTINARIA 120000 2000 11600012000180009000 11900011500017000 1000 2 110008000 Total number of individuals numberTotal individuals of per m 118000 0 11400010000160007000 117000 SH6 1130001500090006000 SH29 120000 11600011200014000 2007 2 50008000 115000119000 1110001300040007000 114000118000 1100001200030006000 113000117000 1090001100020005000 112000116000

1080001000010004000 Total number of individuals numberTotal individuals of per m 111000115000 107000300090000 110000114000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 10600020008000 109000113000 2001 10500010007000 Total number of individuals numberTotal individuals of per m 108000112000 10400060000 107000111000 1030005000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 106000110000 1020004000 2001 105000109000 1010003000 8000108000 1040002000

2 100000 7000103000107000 990001000 6000102000106000 980000 5000101000105000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 97000 100000104000 2001 400096000 10300099000 300095000 10200098000 200094000 10100097000 100093000 Total number of individuals numberTotal individuals of per m 10000096000 920000 9500099000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 91000 9400098000 2007 90000 9300097000 89000 800096000

2 92000 88000 7000 9100095000 87000 6000 9000094000 86000 50008900093000 85000 40008800092000 84000 30008700091000 83000 20008600090000 82000 10008500089000

Total number of individuals numberTotal individuals of per m 81000 84000880000 80000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 8300087000 79000 2012 8200086000 78000 8100085000 Figure 21: Total abundance77000 (no. ind. per m2) of dominant benthic macrofaunal taxonomic groups sampled at 8000084000 eight76000 stations in St Helena Bay in 2001, 2007 and 2012. Data for sites SH31 and SH50 have been 7900083000 omitted75000 from the 2001 adjusted graph to enable the data for the remaining sites to be plotted on the7800082000 same scale as for the other sampling events (2007 and 2012). 74000 7700081000 73000 7600080000 72000 7500079000 Page | 26 71000 7400078000 70000 7300077000 69000 7200076000 68000 7100075000 67000 7000074000 66000 6900073000 65000 6800072000 64000

2 6700071000 63000 6600070000 62000 6500069000 61000 6400068000 2 60000 6300067000 59000 6200066000 58000 6100065000 57000

6000064000

2 Total number of individuals numberTotal individuals of per m 56000 5900063000 55000 5800062000 54000 5700061000

Total number of individuals numberTotal individuals of per m 53000 5600060000 52000 5500059000 51000 5400058000 50000 5300057000 Total number of individuals numberTotal individuals of per m 49000 5200056000 48000 5100055000 47000 5000054000 46000 4900053000 45000 4800052000 44000 4700051000 43000 4600050000 42000 4500049000 41000 4400048000 40000 4300047000 420003900046000 410003800045000 400003700044000 390003600043000 380003500042000 370003400041000 360003300040000 350003200039000 340003100038000 330003000037000 320002900036000 310002800035000 300002700034000 290002600033000 280002500032000 270002400031000 260002300030000 250002200029000 240002100028000 230002000027000 220001900026000 210001800025000 200001700024000 190001600023000 180001500022000 170001400021000 160001300020000 150001200019000 140001100018000 130001000017000 12000160009000 11000150008000 10000140007000 1300090006000 1200080005000 1100070004000 1000060003000 500020009000 400010008000 300070000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 20006000 2001 10005000 40000 3000 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 2000 2001 1000 0 SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 2001

1000 2001 900

800 SIPUNCULIDEA 2 700 POLYCHAETA 600 PENNATULACEA 500 MALACOSTRACA 400 GASTROPODA

300 BRACHYURA Total biomass biomass Total (g) m per 200 BIVALVIA 100 ASCIDIACEA 0 ANTHOZOA SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 2001

1000 900 2007

800 SIPUNCULIDEA 2 700 POLYCHAETA 600 PENNATULACEA 500 MALACOSTRACA 400 GASTROPODA

300 BRACHYURA Total biomass biomass Total (g) m per 200 BIVALVIA 100 ASCIDIACEA 0 ANTHOZOA SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 2007

1000 900 2012

800 SIPUNCULIDEA 2 700 POLYCHAETA 600 PENNATULACEA 500 MALACOSTRACA 400 GASTROPODA

300 BRACHYURA Total biomass biomass Total (g) m per 200 BIVALVIA 100 ASCIDIACEA 0 ANTHOZOA SH6 SH12 SH21 SH26 SH29 SH30 SH31 SH50 2012

Figure 22. Total biomass per m2 of dominant benthic macrofaunal taxonomic groups sampled at eight stations in St Helena Bay in 2001, 2007 and 2012.

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Temporal variation (2007-2012: Inner bay, fish factories and estuary) The average density of macrofauna in St Helena Bay increased between 2007 and 2012 in the bay but decreased in the estuary. The increase in abundance in the bay can be attributed to an increase in polychaetes and malacostraca (notably the amphipods). The benthic macrofaunal community in the bay changed from one dominated by detrivores to a community with a fairly equal proportion of predators, detrivores and filter feeders.

The reduced abundance in the estuary seems to be related to a reduction in polychaete numbers. Malacostraca, notably the mud prawn, Upogebia africana and Callichirus kraussi, increased in abundance in the estuary. The estuarine benthic macrofaunal community shifted from one dominated by detritivores in 2007 to one dominated by filter feeders in 2012.

3000

2500

2 SCAVENGER 2000 PREDATOR

1500 GRAZER

1000

FILTER FEEDER Average abundance abundance Average m per

500 DETRITIVORE

0 2007 2012 2007 2012 Bay Estuary

Figure 23. Average abundance per m2 of benthic macrofauna sampled in the bay and estuary in 2007 and 2012, indicated by functional groups.

3000 POLYCHAETA

2500 OCTOCORALLIA

2 NEMERTEA 2000 MALACOSTRACA

GASTROPODA 1500 CLITELLATA

1000 BRACHYURA

Average abundance abundance Average m per BIVALVIA

500 ACTINARIA

0 2007 2012 2007 2012 Bay Estuary

Figure 24. Average abundance per m2 of benthic macrofauna sampled in the bay and estuary in 2007 and 2012, indicated by taxonomic groups.

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Group average Transform: Fourth root Resemblance: S17 Bray Curtis similarity 0 Year 2007 2012 20

m i

S 60

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4 6 7 9 7 6 8 6 7 6 8 9 3 6 1 0 1 9 0 0 0 1 5 5 4 1 3 6 6 6 9 2 2

5 5 5 4 4 4 4 5 4 4 4 4 5 2 5 3 2 3 5 3 3 5 5 5 2 5 2 2 2 1 1

H H

H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H

S S

S S S S S S S S S S S S S _ S S S S S S S S S S S S S S _ S S S S

______d ______

d d d d d d d d d d d d d d d d

w w w w w w w w w w w w e w w w

l l l l l l l l l l l l l l l l

e e e e e e e e e e e e e e e

O O O O O O O O O O O O O O O O

N N N N N N N N N N N N N N N Samples

Figure 25. Dendrogram indicating similarity among sites in St Helena Bay in 2007 and 2012 based on the benthic macrofaunal community composition. The red lines represent significant clusters based on the SIMPROF analysis.

Transform: Fourth root Resemblance: S17 Bray Curtis similarity Old_SH53 2D Stress: 0.16 Year Old_SH26 Old_SH57 2007 2012 Old_SH31 Old_SH6 Old_SH54 Similarity Old_SH29Old_SH50 20 Old_SH12Old_SH21 Old_SH47 Old_SH30 40 New_SH26New_SH6 Old_SH48 Old_SH55Old_SH56 Old_SH46 New_SH12New_SH29 New_SH55New_SH54 New_SH30 Old_SH49 New_SH21New_SH53New_SH31 New_SH50 New_SH46 New_SH47 New_SH56

New_SH48 New_SH49

Figure 26. Multi-dimension scaling plot representing similarity among sites based on their benthic macrofaunal communities in 2007 and 2012 in St Helena Bay.

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The estuarine sites sampled in 2007 and in 2012 were significantly different to sites sampled in the Bay for both surveys (Figure 25 and Figure 26). Interestingly two of the fish factory sites sampled in 2007 were found to be similar to the estuary sites sampled in 2007, but not significantly so. The SIMPER analysis indicated that the species contributing the most towards the dissimilarity between 2007 and 2012 in the estuary was the detrivorous polychaete Capitella capitata which was found in a high abundance at the estuarine sites in 2007 and was not found in 2012.

Table 5. Species contributing to the differences in the macrofaunal communities in the Berg River estuary in 2007 and 2012. Taxonomic Functional Species Change in abundance Contrib% Cum.% group group Decreased Capitella capitata Detritivore Polychaeta 20.9 20.9 (Absent in 2012) Upogebia africana Filter Feeder Malacostraca Increased 9.86 30.76 Orbinia Increased Detritivore Polychaeta 8.77 39.53 angrapequensis (Absent in 2007) Ceratonereis Decreased Predator Polychaeta 7.62 47.15 erythraeensis (Absent in 2012) Increased Spiroplax spiralis Detritivore Brachyura 5.3 52.45 (Absent in 2007) Increased Nereis spp. Detritivore Polychaeta 5.1 57.55 (Absent in 2007) Callichirus kraussi Detritivore Malacostraca Increased 4.87 62.41 Increased Glycera convoluta Predator Polychaeta 3.46 65.87 (Absent in 2007) Decreased Grandidierella lutosa Detritivore Malacostraca 3.38 69.25 (Absent in 2012)

The results of the ANOSIM test, conducted to compare samples collected in the bay in 2007 to those collected in the bay in 2012, indicated that the benthic macrofaunal communities had changed significantly between 2007 and 2012. The SIMPER analysis revealed that a large range of species contributed to the dissimilarities found between years. The taxa primarily responsible for the dissimilarity were polychaetes and crustaceans. Increases in the abundance of some of the polychaete species (most notably Diopatra monroi and Magelona papillicornis) contributed 24.3% to the overall dissimilarity, while decreases in other polychaete species (most notably Mediomastus capensis and Nephtys sphaerocirrata) contributed 20.9% to the overall dissimilarity between 2007 and 2012. Increases in the abundance of certain crustacean species (most notably Ampelisca spinimana) contributed 18% to the overall dissimilarity, while decreases in other crustacean species contributed 11% to the overall dissimilarity between 2007 and 2012.

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5.2.2 Diversity indices

The diversity of the benthic macrofaunal communities sampled in the bay in 2012 appeared to be very patchy with no clear spatial pattern. The majority of sites sampled had low diversity values (15 sites had an H’ value of less than 1.5).

Figure 27: Diversity (Shannon Weiner Index) of benthic macrofauna interpolated from sites sampled in St Helena Bay in 2012. (H’ = 1.5 indicates low diversity, H’ = 3.5 indicates high diversity)

Temporal variation The majority of sites sampled in 2001 had relatively low diversity values. The diversity at eight of these sites increased between 2001 and 2007 and then decreased again between 2007 and 2012. The estuarine sites along with two of the five fish factory sites had low diversity in 2007, while three of the fish factory sites had a high diversity, and the remainder of sites in the bay had moderately diverse benthic macrofauna communities. The diversity of all of the sites sampled in 2007 was low relative to the 2012 survey with the exception of two estuarine sites, which increased slightly but still had a low diversity value.

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Table 6. Shannon Weiner Diversity Index calculated for sites sampled in St Helena Bay in 2001, 2007 and 2012. Fish factory sites are indicated in bold. (H’ = 1.5 indicates low diversity, H’ = 3.5 indicates high diversity). The changes in diversity between surveys (2001 to 2007 and 2007 to 2012) are indicated in the last two columns with increases highlighted in green and decreases in red.

Site Diversity 2001 Diversity 2007 Diversity 2012 2001 to 2007 2007 to 2012 SH4 1.4 2.0 1.9 Increase Decrease SH6 2.4 2.8 2.0 Increase Decrease SH12 1.9 not sampled 2.0 SH13 1.3 not sampled 2.1 SH14 1.5 not sampled 2.0 SH15 0.7 not sampled 1.7 SH16 not sampled not sampled 1.6 SH21 2.0 2.6 0.9 Increase Decrease SH26 1.4 1.9 0.7 Increase Decrease SH27 2.2 not sampled 2.4 SH29 1.6 2.6 1.4 Increase Decrease SH30 1.7 2.8 1.1 Increase Decrease SH31 0.1 2.5 2.3 Increase Decrease SH33 1.3 not sampled 1.1 SH36 1.0 not sampled 1.1 SH45 not sampled not sampled 1.5 SH46 not sampled 1.0 1.3 Increase SH47 not sampled 0.7 1.3 Increase SH48 not sampled 0.9 0.6 Decrease SH49 not sampled 1.8 0.5 Decrease SH50 1.7 2.5 2.1 Increase Decrease SH51 not sampled not sampled 1.9 SH53 not sampled 2.6 1.0 Decrease SH54 not sampled 1.7 1.1 Decrease SH55 not sampled 2.9 1.6 Decrease SH56 not sampled 3.1 1.0 Decrease SH57 not sampled 1.4 0.0 Decrease

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6 INTEGRATION OF PHYSICO-CHEMICAL PARAMETERS AND BIOTIC INDICATORS

6.1 Methods

The aim of this analysis was to determine how the environmental variables (organic content of sediment, grain size) relate to the observed biological patterns in macrobenthic community structure. This involved superimposing the concentrations of individual environmental variables onto biotic multi-dimensional scaling (MDS) plots. An MDS plot is a spatial representation of the Bray Curtis similarity between sites. MDS plots are constructed using PRIMER V6 from the similarity matrix in order to graphically view similarities between sample sites. Like the dendrogram, samples with similar species composition and abundance cluster together, while those that are less similar are placed further apart. The values of various environmental variables are superimposed on MDS plots as circles of varying diameter (the larger the circle, the higher the concentration). These are known as ‘bubble plots’ and they allow one to easily identify the sites at which contaminants are elevated, as well as to determine if contamination patterns have any correlation to biotic structure.

6.2 Results

The bubble plot based on the 2012 survey results, which incorporated depth values, revealed that the differences between the benthic macrofaunal communities in the estuary and the bay were related to some extent to depth in that all estuarine sites were shallower than the Bay. This is to some extent an artefact of the fact that all sampling sites in the bay were deeper than 4 m and that all of the estuary sites were 1 m depth or less. It may well be that the difference is actually related to difference in salinity or some other parameter not measured in this study. The bubble plots with mud, TOC and TON revealed no clear trend.

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Transform: Fourth root Transform: Fourth root Resemblance: S17 Bray Curtis similarity Resemblance: S17 Bray Curtis similarity 2D Stress: 0.18 Depth 2D Stress: 0.18 Mud SH56 SH56 7 4

SH33 SH33 16 SH51 28 SH51 SH54 SH54 SH47 SH36 SH47 SH36 SH15 SH53 SH15 SH53 49 SH14 28 SH14 SH16 SH16 SH21 SH13SH21 SH13 SH6 SH6 SH46 SH46 SH12 SH12 SH55 SH49 SH55 SH49 70 40

SH27 SH26 SH27 SH26 SH30 SH30 SH48 SH48 SH29 SH29 SH31 SH31SH50 SH50

SH4 SH4

Transform: Fourth root Transform: Fourth root Resemblance: S17 Bray Curtis similarity Resemblance: S17 Bray Curtis similarity 2D Stress: 0.18 2D Stress: 0.18 TOC TON SH56 SH56 0.9 8E-2

SH33 SH33 3.6 SH51 0.32 SH51 SH54 SH54 SH36 SH47 SH36 SH47 SH15 SH53 SH15 SH53 SH14 0.56 SH14 6.3 SH16 SH21 SH16 SH21 SH13 SH13 SH6 SH46 SH6 SH46 SH12 SH12 SH55 SH49 SH55 SH49 9 0.8 SH27 SH27 SH26 SH30 SH26 SH30 SH48 SH48 SH29 SH29 SH31SH50 SH31SH50

SH4 SH4

Figure 28. MDS of St Helena benthic macrofauna abundance (2012) with superimposed circles representing depth, mud, TOC and TON. Circle size is proportional to magnitude of concentration (increasing circle size = larger concentration).

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7 DISCUSSION

The monitoring protocol used in the 2012 survey and previous surveys provides snapshots of the state of physico-chemical environment and the benthic ecology in St Helena Bay since 2001. The positioning of sites allows for discussion on the state of the bay from three different perspectives: the state of the benthic ecosystem in the bay as a whole, the intensity and extent of impacts from point source fish factory effluents, and the state of the lower portion of the Berg River Estuary. The results of the 2012 survey are discussed in the context of these three different perspectives below.

7.1 Bay-wide variability

Sediments Particle size composition of the sediments in the marine environment is influenced by the wave energy, depth and current circulation patterns. Coarser or heavier sand and gravel particles are found in areas with high wave energy and strong currents as the movement of water in these areas suspends fine particles (mud and silt) and flushes these out of the area. Alterations to the wave action and current patterns which reduce or increase the movement of water can result in the deposition (reduced water movement) or removal (increased water movement) of mud from a particular area. The 2012 results indicated a dramatic reduction in the mud content of the sediments in St Helena Bay since 2007 and 2001. This is possibly the result of an increasing intensity and frequency of large storm events, given that the effects have been experienced at a bay-wide scale. Indeed, examinations of storm events over the period 1996 to 2010 by Theron et al. (2010) revealed an increasing trend in the peaks of individual storm events during the stormy winter season offshore of Cape Town.

St Helena Bay is situated downstream of the Cape Columbine upwelling cell. Nutrients are transported equatorward from Cape Columbine into St Helena Bay and retained within a clockwise circulation in the bay (Hutchings et al. 2012). Sun-warming of the surface layers during summer and autumn results in the stratification of the water column which allows for phytoplankton blooms to develop as the cells are kept in the euphotic zone and not mixed deeper within the water column (Bailey 1991). St Helena Bay is an extremely rich zone for plankton due to the enhanced nutrient availability and a stable euphotic zone, and the bay provides a rich feeding ground for planktivores and in turn, predators. At a large scale, the mostly likely source of organic matter in St Helena Bay is from phytoplankton production, the associated detritus that forms from the decay thereof and faecal pellets. Organic matter enrichment from fish factory effluent is evident at a local scale and does not appear to impact the bay as a whole (results of 2012 survey, Carter and Steffani 2007).

The results of the 2012 survey indicated that there had been minor reductions in the levels of organic nitrogen at most sites throughout the bay since 2007, while organic carbon levels decreased in the northern reaches of the bay, increased in the vicinity of the fish factories and underwent minor changes elsewhere in the bay. Interestingly the molar carbon:nitrogen ratios increased dramatically throughout the bay between 2007 and 2012. An increase in the rate of denitrification is likely to have resulted in the reduction of particulate nitrogen in the sediments and the consequent increase in the C:N ratios experienced in St Helena Bay since 2012.

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Denitrification is the process whereby nitrates (organic nitrogen) are reduced by microbes to form nitrogen and ammonia. This process occurs in environments where oxygen levels have been depleted (anoxic or hypoxic) and nitrates are present. In areas where photosynthetic rates are very high, such as in upwelling systems, a high biological oxygen demand deeper in the water column and sediments can lead to complete oxygen utilisation. The highly nutrient enriched, stratified and productive waters of St Helena Bay are conducive to the formation of oxygen deficient conditions (Bailey 1991) and low oxygen events are common (Chapman and Shannon 1985). Denitrifying bacteria are likely to dominate in the area when there is an oxygen deficiency as they are able to substitute the oxygen, required for organic matter degradation, through nitrate reduction and by so doing, become the most efficient energy extractors (Knowles 1982, Tyrrell and Lucas 2002).

The increased rate of denitrification since 2007 may be a result of increased plankton productivity in the bay since 2007 and the consequent reduction in oxygen availability in the sediments. Hutchings et al. (2012) reported decadal scale fluctuations in upwelling-favourable winds observed over the period 1982 to 2006, with winds reaching a peak in 2001 and subsequently undergoing a decline to 2006. El Niño events have at times been known to suppress upwelling along the coast, while La Niña increases upwelling intensity (Arntz et al. 2006, Rouault et al. 2010). The 2012 survey was conducted following several years in which La Niña events dominated while the 2007 survey was conducted following several years in which El Niño events dominated. This could be an indication that upwelling-favourable conditions and therefore plankton productivity were higher in years preceding the 2012 survey compared to years preceding the 2007 survey. However it is important to note that not all events are experienced by the Benguela system as they are in the Pacific systems (Arntz et al. 2006).

20 La Niña

-10

-20 El Niño Southern OscillationIndex Southern

-30

1996 1997 1998 1999 2000 2005 2006 2007 2008 2009 2001 2002 2003 2004 2010 2011 2012

Figure 29. Time series of the Southern Oscillation Index, indicating El Niño and La Niña events since 1996 and highlighting years in which St Helena Bay has been surveyed. Note the El Niño event of 1997/1998 was not experienced in the Benguela system. (Data source: Arntz et al. 2006 and http://www.bom.gov.au/climate/glossary/soi.shtml).

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Benthic macrofauna Changes in benthic species composition can be the first indicator of disturbance or habitat alteration, as certain species are more sensitive (i.e. likely to decrease in abundance in response to stress) while others are more tolerant of adverse conditions (and may increase in abundance in response to stress, taking up space or resources vacated by the more sensitive species). Certain species are able to rapidly invade and colonise disturbed areas as they have high fecundity, rapid growth and rather short life-cycles (Newell 1998). These species are known as “r-strategists” or opportunistic species and their presence generally indicates unpredictable short-term variations in environmental conditions which may result from either natural factors or anthropogenic activities. In stable environments the community composition is controlled predominantly by biological interactions rather than by fluctuations in environmental conditions. Species found in these conditions are known as “K-strategists” and are selected for their competitive ability. K-strategists are characterised by long-life spans, larger body sizes, delayed reproduction and low death rates. Intermediate communities with different relative proportions of opportunistic species and K- strategists are likely to exist between the extremes of stable and unstable environments.

Eastern Boundary Current ecosystems, such as the BCS are generally known to be dominated by opportunistic benthic species with short life spans and relatively high reproductive rates (r- strategists), which are adapted to surviving in oxygen deficient conditions (Arntz et al. 2006). Such opportunistic species should recover rapidly following natural or anthropogenic disturbances. The 2012 survey revealed that the benthic macrofaunal community in St Helena Bay is dominated by the predatory polychaete Diopatra monroi both in terms of abundance and biomass. The filter feeding amphipod Ampelisca anomala, also dominated in terms of abundance and the clam, Venerupis corrugata, in terms of biomass. These species are typical of the continental shelf sediments of the west coast (Christie 1976, Carter and Steffani 2006, Carter and Steffani 2007). The dominance of amphipods suggests that the system has a fairly low level of contamination as amphipods have a lower tolerance than other groups to pollution, particularly organic pollution (Christie and Moldan 1977).

Polychaetes and amphipods, which have dominated the benthic macrofauna in terms of abundance since 2001, decreased between 2001 and 2007 and then increased between 2007 and 2012. Similarly bivalves and polychaetes, which have dominated in terms of biomass since 2001, underwent a reduction between 2001 and 2007 and then an increase between 2007 and 2012. Diversity values increased between 2001 and 2007 and then decreased between 2007 and 2012. This suggests that the benthic communities were subjected to a bay-wide variation or disturbance, potentially of a repetitive nature, and that at lower densities the community is able to support greater species diversity. As suggested above for the changes in the C:N ratios, the productivity of phytoplankton and the consequent levels of oxygen availability may have varied in the bay, possibly due to decadal scale fluctuations in upwelling-favourable winds. The change detected in C:N ratios between 2007 and 2012 and the known occurrence of denitrification in the BCS, suggests that oxygen availability in the sediments was higher in 2007 than in 2012. This is likely to have had consequences for the benthic macrofauna community structure. It may have favoured environmental conditions that enabled species to share habitat and outcompete those opportunistic species usually dominating in oxygen deficient conditions.

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The sea pen (Virgularia schultzei) is typical of the west coast, but unlike other west coast species, is relatively slow growing and is thus more sensitive to variation. The sea pen was present at two sites in 2001 at relatively low densities and was not found again in 2007 and 2012. This suggests that the benthic ecosystem underwent a disturbance to the state that is usually suited to sea pens.

The results of 2012 survey further emphasized the high natural variability of benthic soft-bottom communities. The benthic communities in the bay were not spatially grouped and did not clearly correlate with any of the environmental variables measured (depth, TOC, TON, mud content). It is likely that the communities in the bay are influenced by complex biological interactions at the sediment-water interface, various physico-chemical properties and potentially an interplay between those properties and biological interactions.

7.2 Fish factory sites

Given the known impacts of fish factory discharges on the marine environment (Christie and Moldan 1977) the areas around the fish factory discharge point were prioritised as key sites for monitoring purposes in the 2007 and 2012 surveys. Results of the 2012 survey revealed that the areas in the vicinity of the fish factory outfalls were carbon enriched compared to other areas in the bay and that the levels of enrichment at these sites had increased disproportionately compared to other sites in the bay. This suggests that the effluent from fish factories is having a marked effect on the organic composition of the sediments at a local scale (sites within 50m of fish factory outfalls) but not at a large scale.

The overlapped effects of anthropogenic and natural disturbances on benthic communities present a challenge in environmental management and in some cases the health of an ecosystem becomes difficult to evaluate when natural disturbances mask the response of potential ecological indicators. For this reason it is important to assess the data from specific sites in the context of the larger system. However, all of the sites sampled adjacent to the fish factory effluent pipes were found to be significantly different from one another. Two sites sampled in the immediate vicinity of the fish factory discharge points were in fact outliers and were characterised by depauperate benthic macrofaunal communities. No benthic macrofauna were found in the sample taken from SH57 and only three polychaete species, with very low abundances were sampled at SH 56. Given that the macrofauna on the west coast are typically opportunistic species, able to recover relatively rapidly from disturbances and that the communities at site SH56 and SH57 were in such a poor state, it is evident that effluents from these fish factories are having a negative impact upon the benthic macrofaunal community at these sites. However, as the results of the 2007 survey indicated, these impacts are limited to the local area around the fish factories discharge points.

7.3 Estuarine sites

The levels of carbon and nitrogen enrichment in the estuary remained relatively low between 2007 and 2012, however the C:N rations increased substantially, possibly as a combined result of terrigenous nutrient inputs and nutrient inputs from the marine environment. The benthic

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macrofaunal communities sampled in the Berg River Estuary were significantly different to those sampled in the Bay and were characterised by the presence of the estuarine mud prawn (Upogebia africana), two detritivorous polychaetes species (Orbinia angrapequensis and Nereis spp.) and the three-legged crab (Spiroplax spiralis). Changes since the 2007 survey include a reduction in the overall abundance of macrofaunal abundance, particularly polychaetes, an increase in prawn abundance and a shift in community structure from a system dominated by detritivores to one dominated by filter feeders.

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8 CONCLUSIONS

 Both sediment results (C:N ratios) and benthic community results indicate that bay-wide fluctuations are occurring. These are possibly linked to decadal-scale variations in upwelling intensity and phytoplankton productivity;  Monitoring in the larger bay area must continue to aid in the interpretation of variability detected at local scales;  The levels of organic contamination have increased at two of the sites adjacent to fish factories (fish factory in Stompneusbaai and the most southerly fish factory) as well as in Britannia Bay since 2007;  The benthic macrofauna sampled at the two heavily contaminated fish factory sites provide evidence of a negative impact to the marine ecology at a local scale; and  There is evidence of organic enrichment at the site situated in Britannia Bay. Sources of this enrichment should be investigated.

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9 REFERENCES

Arntz, W.E., Gallardo, V.A., Gutierrrez, D., Isla, E., Levin, L.A., Mendo, J., Neira, C., Rowe, G.T., Tarazona, J. and M. Wolff. 2006. El Nino and similar perturbation effects on the benthos of the Humboldt, California, and Benguela Current upwelling ecosystems. Advances in Geosciences, 6, 243-265. Bailey, G.W. 1991. Organic carbon flux and development of oxygen deficiency on the modern Benguela continental shelf south of 22°S: spatial and temporal variability. Modern and Ancient Continental Shelf Anoxia. Geological Society Special Publication No. 58, 171-183 Bakun, A. 1998. Ocean triads and radical interdecadal stock variability: bane and boon for fishery management science. In: Pitcher, T.J., Hart, P.J.B., Pauly, D. (Eds.), Reinventing Fisheries Management. Kluwer Academic Publishers, Dordrecht, Netherlands, pp. 331–358.Bakun 1998, Branch, G. M., & Branch, M. 1981. The Living Shores of Southern Africa. C. Struik, Cape Town. Carter, R. and N. Steffani. 2007. State of Saint Helena Bay 2007 Benthic Macrofauna Distributions. Prepred for CSIR. Chapman, P. & L.V. Shannon. 1985. The Benguela Ecosystem. 2 Chemistry and related processes. Oceanography and Marine Biology Annual Review 23, 183-251. Christie, N.D. 1976. A numerical analysis of the distribution of a shallow sublittoral sand macrofauna along a transect at Lamberts Bay, South Africa. Trans. R. Soc. S. Afr. 42: 149-Christie 1976 Christie, N.D. & A. Moldan. 1977. Effects of Fish Factory Effluent on the Benthic Macrofauna of Saldanha Bay. Marine Pollution Bulletin, 8, 2, 41-45. Cockcroft, A.C., Schoeman, D.S., Pitcher, G.C., Bailey, G.W. & D.C. Van Zyl 2000 -A mass stranding, or "walkout" of west coast rock lobster Jasus lalandii in Elands Bay, South Africa: causes, results and implications. In The Biodiversity Crises and Crustacea.Von Kaupel Klein, J.C. and F.R. Schram (Eds). Crustacean Iss. 11: 362-688. CSIR 2008. St Helena Bay Water Quality Trust: The Biogeochemical status of surface sediments in St Helena Bay in 2007. Prepared for the St Helena Bay Water Quality Trust Hutchings, L., Jarre, A., Lamont, T. & M. Van den Berg. In Press. St Helena Bay, then and now: muted climate signals, large human impact. African Journal of Marine Science. Knowles, R. 1982. Denitrification. Microbiological Reviews 46 (1), 43-70 Monteiro, P.M.S. & A.M. Roychoudhury. 2005. Spatial Characteristics of sediment trace metals in an eastern boundary upwelling retention area (St. Helena Bay, South Africa): A hydrodynamic- biological pump hypothesis. Estuarine, Coastal and Shelf Science, 65, 123-134. Rouault, M., Pohl, B. & P. Penven. 2010. Coastal oceanic climate change and variability from 1982 to 2009 around South Africa. African Journal of Marine Science.32(2):237-246 Shannon, L.V. & M.J. O’Toole, 1998. BCLME Thematic Report 2: Integrated overview of the oceanography and environmental variability of the Benguela Current region. Unpublished BCLME Report, 58pp Theron, A. , M.Rossouw , L. Barwell , A.Maherry , G.Diedericks & P. de Wet. 2010. Quantification of risks to coastal areas and development: wave run-up and erosion. NE20-PA-F Tyrrell, T. & M. Lucas. 2002. Geochemical evidence of denitrification in the Benguela upwelling system. Continental Shelf Research, 22, 2497-2511 Warwick, R.M. 1993. Environmental impact studies on marine communities: Pragmatical considerations. Australian Journal of Ecology 18: 63-80.

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ANNEXURE 1: SEDIMENT PARTICLE SIZE, TOC AND TON MEASURED IN 2012

Site TOC TON Sand Mud SH 4 2.976 0.299 92.45 7.55 SH 6 0.269 0.016 99.84 0.16 SH 12 4.355 0.456 90.91 9.09 SH 13 3.744 0.409 89.75 10.25 SH 14 3.612 0.357 95.48 4.52 SH 15 3.885 0.401 96.14 3.86 SH 16 2.592 0.272 82.84 17.16 SH 21 0.597 0.04 94.33 5.67 SH 26 4.496 0.439 97.59 2.41 SH 27 1.069 0.081 98.94 1.06 SH 29 0.17 0.005 98.41 1.59 SH 30 0.324 0.018 99.62 0.38 SH 31 0.229 0.011 99.19 0.81 SH 33 3.244 0.321 67.95 32.05 SH 36 1.856 0.178 93.06 6.94 SH 46 0.902 0.073 83.93 16.07 SH 47 0.308 0.012 99.35 0.65 SH 48 0.54 0.035 96.47 3.53 SH 49 0.434 0.025 97.32 2.68 SH 50 0.446 0.032 84.77 15.23 SH 51 8.541 0.429 99.30 0.70 SH 53 7.158 0.711 98.54 1.46 SH 54 7.108 0.596 94.20 5.80 SH 55 2.557 0.152 83.57 16.43 SH 56 3.589 0.08 99.88 0.12 SH 57 3.78 0.354 96.51 3.49

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ANNEXURE 2: BENTHIC MACROFAUNA SAMPLED IN 2012 (abundance per m2)

SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH Afrophaxas decipiens 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 6.9 0.0 0.0 Amaryllis macrophthalma 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ampelisca anomola 0.0 0.0 0.0 2571.2 4225.2 472.6 0.0 4774.1 0.0 0.0 0.0 27.8 0.0 708.8 8394.7 139.0 0.0 20.8 0.0 0.0 6.9 0.0 34.7 0.0 0.0 0.0 0.0 Ampelisca brevicornis 0.0 69.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ampelisca palmata 0.0 20.8 0.0 2800.6 708.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 48.6 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ampelisca spinimana 0.0 0.0 840.9 0.0 0.0 0.0 173.7 0.0 13.9 236.3 6.9 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 6.9 0.0 0.0 952.1 0.0 0.0 Amphieteis gunneri 0.0 0.0 0.0 62.5 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Amphilochus neapolitanus 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Anemone 0.0 13.9 312.7 229.3 132.0 104.2 118.1 13.9 0.0 69.5 722.7 145.9 111.2 0.0 97.3 132.0 6.9 0.0 0.0 0.0 0.0 132.0 173.7 27.8 76.4 0.0 0.0

Anemone 2 6.9 0.0 0.0 6.9 0.0 0.0 0.0 13.9 0.0 0.0 27.8 69.5 83.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 62.5 0.0 0.0 0.0 0.0 0.0 0.0

Anemone 3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Anthelura remipes 6.9 0.0 0.0 6.9 55.6 20.8 0.0 48.6 0.0 0.0 208.5 104.2 83.4 0.0 34.7 41.7 0.0 0.0 6.9 0.0 48.6 0.0 0.0 0.0 6.9 0.0 0.0

Arenicola loveni 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 250.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0

Ascidian 0.0 20.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Bathyporeia sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0

Bullia laevissima 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0

Callichirus kraussi 0.0 0.0 0.0 6.9 6.9 41.7 118.1 104.2 0.0 0.0 0.0 6.9 0.0 0.0 0.0 13.9 0.0 166.8 0.0 0.0 0.0 0.0 34.7 0.0 6.9 0.0 0.0 Calyptraea chinensis 0.0 0.0 0.0 465.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Capitella capitata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 514.2 0.0 0.0 0.0

Caprellid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Caprellid 2 0.0 0.0 0.0 0.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH Choromytilus meridionalis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 20.8 0.0 0.0 0.0 0.0

Cirolana hirtipes 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Cirolana sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Cirriformia tentaculata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0

Corophiid 1 0.0 90.3 145.9 549.0 1855.5 2293.3 145.9 0.0 20.8 0.0 0.0 0.0 0.0 90.3 76.4 500.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Corophiid 2 0.0 0.0 0.0 0.0 0.0 0.0 27.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0

Crepidula spp. 0.0 0.0 0.0 20.8 0.0 41.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 27.8 0.0 0.0 0.0 0.0 0.0 0.0 118.1 0.0 0.0 0.0 0.0

Cumacea sp. 2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 20.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Diopatra monroi 0.0 889.5 257.1 465.6 3182.8 2911.7 1820.7 840.9 535.1 694.9 62.5 13.9 0.0 0.0 0.0 4725.5 0.0 0.0 0.0 0.0 0.0 347.5 4975.7 625.4 62.5 0.0 0.0

Discinisca tenuis 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Dosinia lupinus orbignyi 6.9 34.7 13.9 76.4 13.9 6.9 6.9 0.0 83.4 542.0 6.9 0.0 0.0 0.0 27.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Dromidia sp. 0.0 41.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Eunicella papillosa 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Fusciolariidae 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Gammaropsis sophiae 0.0 0.0 361.4 20.8 0.0 0.0 458.7 0.0 0.0 125.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Glycera convoluta 0.0 41.7 0.0 0.0 13.9 6.9 0.0 0.0 0.0 27.8 0.0 6.9 13.9 0.0 0.0 0.0 6.9 6.9 0.0 0.0 0.0 0.0 20.8 6.9 6.9 0.0 0.0

Harmothoe spp. 0.0 0.0 6.9 27.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Hippomedon onconotus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 41.7 0.0 0.0 0.0 0.0 0.0 Hymensoma obiculare 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Iphinae sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 152.9 0.0 97.3 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 48.6 0.0 0.0 0.0 0.0 0.0 0.0 Lemboides crenatipalma 0.0 6.9 83.4 145.9 576.8 632.4 48.6 0.0 0.0 0.0 0.0 0.0 0.0 20.8 13.9 236.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Leucothoe richiardi 0.0 20.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Listriella lindae 0.0 0.0 6.9 0.0 41.7 76.4 20.8 27.8 0.0 6.9 0.0 0.0 0.0 34.7 41.7 27.8 0.0 0.0 0.0 0.0 0.0 0.0 20.8 0.0 13.9 0.0 0.0 Lumbrinereis heteropoda 20.8 6.9 27.8 83.4 41.7 13.9 34.7 27.8 6.9 27.8 55.6 20.8 0.0 0.0 6.9 69.5 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH difficilis Lysianassa ceratina 0.0 48.6 0.0 13.9 0.0 0.0 0.0 0.0 0.0 20.8 0.0 0.0 0.0 0.0 41.7 0.0 41.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Macoma crawfordi ordinaria 6.9 76.4 0.0 145.9 194.6 145.9 76.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 180.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 27.8 41.7 0.0 0.0

Maera inaequipes 0.0 6.9 0.0 132.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Magelona papillicornis 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 13.9 5017.4 368.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 222.4 0.0 0.0 0.0 20.8 0.0 0.0

Melita sp. 0.0 0.0 0.0 6.9 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0

Melita subchelata 0.0 0.0 27.8 76.4 701.9 83.4 76.4 41.7 0.0 6.9 0.0 0.0 0.0 48.6 1042.4 76.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nassarius vinctus 27.8 27.8 0.0 41.7 90.3 180.7 0.0 0.0 0.0 201.5 0.0 0.0 0.0 0.0 48.6 97.3 0.0 0.0 0.0 0.0 0.0 0.0 104.2 0.0 0.0 0.0 0.0

Nebalia capensis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nematode 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Nephtys hombergii 0.0 0.0 6.9 0.0 13.9 0.0 0.0 27.8 0.0 0.0 0.0 333.6 166.8 0.0 20.8 6.9 0.0 13.9 0.0 0.0 125.1 111.2 13.9 13.9 69.5 6.9 0.0 Nephtys sphaerocirrata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 34.7 0.0 0.0 0.0 0.0 111.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 27.8 0.0 0.0

Nereis spp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 6.9 0.0 13.9 0.0 20.8 0.0 0.0 20.8 0.0 0.0 0.0 0.0 0.0 Notomastus latericeus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nucula nucleus 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Orbinia angrapequensis 0.0 0.0 0.0 159.8 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 6.9 27.8 6.9 6.9 0.0 0.0 0.0 0.0 13.9 0.0 0.0

Owenia fusiformis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 180.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Paramoera capensis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 597.6 0.0 0.0 0.0 0.0 0.0 Paraprionospio pinnata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Pectinaria capensis 0.0 0.0 0.0 27.8 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 271.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 486.4 0.0 41.7 0.0 208.5 13.9 0.0 Pherusa swakopiana 0.0 6.9 222.4 0.0 0.0 0.0 0.0 0.0 0.0 166.8 76.4 34.7 48.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 Photis longidactylus 0.0 20.8 0.0 0.0 173.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 785.3 0.0 0.0 0.0 0.0 0.0 0.0 333.6 0.0 0.0 0.0 0.0 0.0

Photis longimanus 0.0 6.9 0.0 6.9 132.0 90.3 0.0 111.2 0.0 0.0 0.0 0.0 0.0 159.8 820.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH Pseudopotamilla reniformis 0.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Pterygosquilla armata capensis 0.0 6.9 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Pycnogonid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Sabellides luderitzi 0.0 0.0 0.0 0.0 20.8 0.0 0.0 0.0 0.0 13.9 13.9 1237.0 646.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 97.3 0.0 0.0 Schistomeringos rudolphii 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Sipunculid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 20.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Sphaerodorum gracile 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0

Sphaeromatidae 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Spiroplax spiralis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 34.7 0.0 13.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Tellimya sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 222.4 0.0 0.0 0.0 0.0 0.0 0.0

Tellina gilchristi 6.9 104.2 312.7 180.7 278.0 0.0 0.0 0.0 6.9 152.9 0.0 0.0 0.0 0.0 6.9 562.9 0.0 0.0 0.0 0.0 0.0 20.8 0.0 0.0 0.0 0.0 0.0

Tellina trilatera 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 104.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Upogebia capensis 0.0 0.0 0.0 0.0 41.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0

Upogebia africana 0.0 0.0 0.0 6.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 0.0 0.0 0.0 152.9 0.0 180.7 291.9 250.2 0.0 0.0 0.0 0.0 0.0 0.0

Urothoe grimaldi 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 34.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 333.6 159.8 0.0 0.0 0.0 0.0 0.0 Venerupis corrugatus 0.0 0.0 0.0 298.8 173.7 83.4 48.6 34.7 0.0 0.0 0.0 0.0 0.0 0.0 6.9 736.6 6.9 0.0 0.0 0.0 0.0 20.8 729.7 20.8 0.0 0.0 0.0 Virgularia schultzei 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 145.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 13.9 0.0 222.4 0.0 0.0 0.0 0.0

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ANNEXURE 3: BENTHIC MACROFAUNA SAMPLED IN 2012 (biomass per m2)

SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH4 Afrophaxas decipiens 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 26.7 0.0 0.0 2.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 Amaryllis macrophthalma 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ampelisca anomola 0.0 0.0 0.0 10.7 10.9 1.2 0.0 26.6 0.0 0.0 0.0 0.1 0.0 3.1 25.3 0.3 0.0 0.1 0.0 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 Ampelisca brevicornis 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Ampelisca palmata 0.0 0.0 0.0 10.5 3.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ampelisca spinimana 0.0 0.0 2.2 0.0 0.0 0.0 0.6 0.0 0.0 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4.1 0.0 0.0

Amphieteis gunneri 0.0 0.0 0.0 9.2 0.0 0.0 1.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Amphilochus neapolitanus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Anemone 0.0 4.8 64.8 36.5 24.0 19.5 50.4 0.3 0.0 2.8 99.0 26.6 8.2 0.0 33.7 30.7 0.1 0.0 0.0 0.0 0.0 25.0 36.9 5.2 80.1 0.0 0.0

Anemone 2 44.6 0.0 0.0 80.5 0.0 0.0 0.0 45.8 0.0 0.0 39.9 31.9 33.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 23.3 0.0 0.0 0.0 0.0 0.0 0.0

Anemone 3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Anthelura remipes 0.3 0.0 0.0 0.1 0.6 0.7 0.0 0.6 0.0 0.0 3.1 2.4 2.2 0.0 0.9 0.8 0.0 0.0 0.1 0.0 1.6 0.0 0.0 0.0 0.3 0.0 0.0

Arenicola loveni 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 630.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 27.3 0.0 0.0

Ascidian 0.0 106.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Bathyporeia sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Bullia laevissima 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 141.7 0.0 0.0 0.0 0.0 0.0 41.4 0.0 0.0 0.0 0.0 0.0 0.0

Callichirus kraussi 0.0 0.0 0.0 0.8 1.2 11.3 41.8 6.9 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.8 0.0 74.7 0.0 0.0 0.0 0.0 0.4 0.0 1.0 0.0 0.0 Calyptraea chinensis 0.0 0.0 0.0 94.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Capitella capitata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.8 0.0 0.0 0.0

Caprellid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Caprellid 2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Choromytilus meridionalis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 839.2 435.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH4

Cirolana hirtipes 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Cirolana sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Cirriformia tentaculata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3.1 0.0

Corophiid 1 0.0 0.1 0.8 1.3 5.8 8.0 0.8 0.0 0.1 0.0 0.0 0.0 0.0 0.3 0.4 1.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Corophiid 2 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Crepidula spp. 0.0 0.0 0.0 3.3 0.0 4.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3.0 0.0 0.0 0.0 0.0 0.0 0.0 51.8 0.0 0.0 0.0 0.0

Cumacea sp. 2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Diopatra monroi 0.0 145.5 68.2 116.6 281.5 323.3 291.9 226.2 102.7 114.1 27.1 7.4 0.0 0.0 0.0 574.3 0.0 0.0 0.0 0.0 0.0 7.4 2207.2 120.3 39.1 0.0 0.0

Discinisca tenuis 0.0 0.0 0.0 27.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Dosinia lupinus orbignyi 29.1 89.8 72.5 443.2 66.5 50.9 66.8 0.0 173.3 539.5 0.0 0.0 0.0 0.0 95.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Dromidia sp. 0.0 97.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Eunicella papillosa 0.0 19.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Fusciolariidae 0.0 0.0 0.0 0.0 76.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Gammaropsis sophiae 0.0 0.0 0.2 0.0 0.0 0.0 0.3 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Glycera convoluta 0.0 3.9 0.0 0.0 6.1 6.0 0.0 0.0 0.0 26.3 0.0 7.8 0.1 0.0 0.0 0.0 0.3 8.8 0.0 0.0 0.0 0.0 3.5 0.1 5.1 0.0 0.0

Harmothoe spp. 0.0 0.0 0.2 1.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Hippomedon onconotus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 Hymensoma obiculare 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 14.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Iphinae sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.7 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 Lemboides crenatipalma 0.0 0.0 0.7 0.8 2.2 2.6 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Leucothoe richiardi 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Listriella lindae 0.0 0.0 0.0 0.0 0.5 1.1 0.5 0.2 0.0 0.1 0.0 0.0 0.0 0.5 0.7 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.2 0.0 0.0 Lumbrinereis heteropoda difficilis 30.8 14.1 41.8 131.9 90.2 19.0 74.5 55.7 7.8 20.8 67.1 45.2 0.0 0.0 19.7 112.3 0.0 0.0 0.0 0.0 1.6 0.0 0.0 0.0 0.0 0.0 0.0 Lysianassa ceratina 0.0 0.8 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.5 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH4 Macoma crawfordi ordinaria 1.9 69.8 0.0 199.1 152.3 137.5 94.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 67.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 10.6 1.1 0.0 0.0

Maera inaequipes 0.0 0.2 0.0 2.2 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Magelona papillicornis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 27.3 0.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.7 0.0 0.0 0.0 0.4 0.0 0.0

Melita sp. 0.0 0.0 0.0 0.0 0.0 0.1 0.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0

Melita subchelata 0.0 0.0 0.2 0.3 3.2 0.5 0.0 0.2 0.0 0.1 0.0 0.0 0.0 0.4 5.5 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nassarius vinctus 9.5 6.3 0.0 11.0 33.8 74.1 0.0 0.0 0.0 68.9 0.0 0.0 0.0 0.0 9.1 30.4 0.0 0.0 0.0 0.0 0.0 0.0 32.5 0.0 0.0 0.0 0.0

Nebalia capensis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nematode 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nephtys hombergii 0.0 0.0 3.9 0.0 3.2 0.0 0.0 9.7 0.0 0.0 0.0 5.1 9.4 0.0 9.8 3.7 0.0 20.2 0.0 0.0 38.4 64.0 7.7 0.4 33.4 0.1 0.0 Nephtys sphaerocirrata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0

Nereis spp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 1.9 0.0 0.1 0.0 0.1 0.0 0.0 0.6 0.0 0.0 0.0 0.0 0.0 Notomastus latericeus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Nucula nucleus 0.0 6.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Orbinia angrapequensis 0.0 0.0 0.0 41.5 1.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.1 1.7 0.3 0.1 0.0 0.0 0.0 0.0 0.6 0.0 0.0

Owenia fusiformis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Paramoera capensis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1.7 0.0 0.0 0.0 0.0 0.0 Paraprionospio pinnata 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Pectinaria capensis 0.0 0.0 0.0 4.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 10.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 58.1 0.0 83.8 0.0 48.3 0.1 0.0 Pherusa swakopiana 0.0 3.7 133.4 0.0 0.0 0.0 0.0 0.0 0.0 27.3 52.1 31.5 78.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3.5 0.0 0.0 Photis longidactylus 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.5 0.0 0.0 0.0 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.0 0.0

Photis longimanus 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.1 0.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Pseudopotamilla reniformis 0.0 5.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Pterygosquilla armata capensis 0.0 17.7 0.0 0.0 0.0 15.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 10.4 12.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Pycnogonid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

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SH6 SH12 SH13 SH14 SH15 SH16 SH21 SH26 SH27 SH29 SH30 SH31 SH33 SH36 SH45 SH46 SH47 SH48 SH49 SH50 SH51 SH53 SH54 SH55 SH56 SH57 SH4

Sabellides luderitzi 0.0 0.0 0.0 0.0 3.9 0.0 0.0 0.0 0.0 0.1 0.2 15.8 18.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2.5 0.0 0.0 Schistomeringos rudolphii 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Sipunculid 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 19.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Sphaerodorum gracile 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0

Sphaeromatidae 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.6 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Spiroplax spiralis 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2.5 0.0 1.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Tellimya sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.2 0.0 0.0 0.0 0.0 0.0 0.0

Tellina gilchristi 6.2 46.1 440.4 125.9 105.3 0.0 0.0 0.0 4.7 120.8 0.0 0.0 0.0 0.0 0.9 286.8 0.0 0.0 0.0 0.0 0.0 11.6 0.0 0.0 0.0 0.0 0.0

Tellina trilatera 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 113.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

Upogebia capensis 0.0 0.0 0.0 0.0 4.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.7 0.0 0.0 0.0 0.0

Upogebia africana 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.4 0.0 0.0 0.0 0.0 88.0 0.0 136.5 275.2 16.0 0.0 0.0 0.0 0.0 0.0 0.0

Urothoe grimaldi 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.9 0.4 0.0 0.0 0.0 0.0 0.0 Venerupis corrugatus 0.0 0.0 0.0 823.0 822.8 439.8 124.0 325.6 0.0 0.0 0.0 0.0 0.0 0.0 18.3 3712.7 5.5 0.0 0.0 0.0 0.0 3.4 2773.4 1.5 0.0 0.0 0.0

Virgularia schultzei 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 34.5 96.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2.8 0.0 1545.4 0.0 0.0 0.0 0.0

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