St Helena Bay State of the Bay 2012 - Anchor Environmental

ST HELENA BAY WATER

QUALITY TRUST:

St Helena Bay

State of the Bay 2012

Prepared by

ANCHORe n v i r o n m e n t a l

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:

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

ANCHORe n v i r o n m e n t a l

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Executive summary

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, one of four major eastern-boundary current systems which is characterised by the

wind-driven upwelling of cold, nutrient rich water. 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. The bay is 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. The Bay also serves as a major node for industrial and small-scale

fisheries on the west coast, as well as for other industries such as mariculture, ship repair and

shipbuilding.

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.

Lat (S) Lon (E) Site Name

2001 2007 2012

Macrofauna Sediment Macrofauna Sediment Macrofauna and

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


32.7078 18.0240 SH12 X X X X X





32.7250 18.1238 SH21 X X X X X

32.4245 18.1248 SH26 X X X 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





32.7000 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




32.6968 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





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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.

Site Depth (m) Gravel (%) Sand (%) Mud (%) TOC

(ppm) TON

(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).

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

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

SH 4

SH 6

SH 1

2

SH 1

3

SH 1

4

SH 1

5

SH 1

6

SH 2

1

SH 2

6

SH 2

7

SH 2

9

SH 3

0

SH 3

1

SH 3

3

SH 3

6

SH 4

6

SH 4

7

SH 4

8

SH 4

9

SH 5

0

SH 5

1

SH 5

3

SH 5

4

SH 5

5

SH 5

6

SH 5

7

Mud

Sand

Gravel

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

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

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

SH 4

SH 6

SH 1

2

SH 1

3

SH 1

4

SH 1

5

SH 1

6

SH 2

1

SH 2

6

SH 2

7

SH 2

9

SH 3

0

SH 3

1

SH 3

3

SH 3

6

SH 4

6

SH 4

7

SH 4

8

SH 4

9

SH 5

0

SH 5

1

SH 5

3

SH 5

4

SH 5

5

SH 5

6

SH 5

7

Mud

Sand

Gravel

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

SH 4

SH 6

SH 1

2

SH 1

3

SH 1

4

SH 1

5

SH 1

6

SH 2

1

SH 2

6

SH 2

7

SH 2

9

SH 3

0

SH 3

1

SH 3

3

SH 3

6

SH 4

6

SH 4

7

SH 4

8

SH 4

9

SH 5

0

SH 5

1

SH 5

3

SH 5

4

SH 5

5

SH 5

6

SH 5

7

Mud

Sand

Gravel

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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.

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

0.00

1.00

2.00

3.00

4.00

5.00

6.00

7.00

8.00

9.00

SH 4

SH 6

SH 1

2

SH 1

3

SH 1

4

SH 1

5

SH 1

6

SH 2

1

SH 2

6

SH 2

7

SH 2

9

SH 3

0

SH 3

1

SH 3

3

SH 3

6

SH 4

6

SH 4

7

SH 4

8

SH 4

9

SH 5

0

SH 5

1

SH 5

3

SH 5

4

SH 5

5

SH 5

6

SH 5

7

TOC

(p

pm

)

2007

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.

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

0.00

0.20

0.40

0.60

0.80

1.00

1.20

1.40

SH 4

SH 6

SH 1

2

SH 1

3

SH 1

4

SH 1

5

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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.

Page | 14

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

Fish Factories

SH 53 9.2 11.7

SH 54 7.8 13.9

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

Southern reaches

SH 12 8.2 11.1

SH 13 7.9 10.7

SH 14 7.9 11.8

SH 15 8.1 11.3

SH 16 7.7 11.1

SH 21 6.7 17.4

SH 33 7.8 11.8

SH 36 7.9 12.2

Eastern inshore

SH 29 6.6 39.7

SH 30 6.0 21.0

SH 31 4.7 24.3

SH 50 5.3 16.3

Northern Reaches SH 26 9.4 11.9

SH 27 9.3 15.4

Berg River Estuary

SH 46 6.9 14.4

SH 47 6.6 29.9

SH 48 7.2 18.0

SH 49 7.0 20.3

Page | 15

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

Page | 16

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

Page | 17

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.

Page | 18

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.

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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SIPUNCULIDEA

PYCNOGONIDA

POLYCHAETA

MALACOSTRACA (ISOPODA)

MALACOSTRACA (BRACHYURA)

MALACOSTRACA (AMPHIPODA)

MALACOSTRACA

GASTROPODA

BRACHIOPODA

BIVALVIA

ASCIDIACEA

ANTHOZOA

Page | 19

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

Taxonomic Level

% Abundance % Biomass Phylum Class Species

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).

Page | 20

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.

0%

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SCAVENGER

PREDATOR

GRAZER

FILTER FEEDER

DETRITIVORE

Page | 21

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.

A

BC

D

E

Page | 22

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

Bay in 2012.

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) Photograph by: N. Steffani

Glycera convoluta (Predator) Photograph by: N. Steffani

Page | 24

MALACOSTRACA (Amphipoda)

Hippomedon normalis (Scavenger) Photograph by: N. Steffani

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

MALACOSTRACA (Decapoda)

Callichirus kraussi (Detritivore) Photograph by: C. Griffiths

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

GASTROPODA

Nassarius vinctus (Scavenger)

Photograph by: N. Steffani

Venerupis corrugata (Filter feeder)

Page | 25

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 | 26

Figure 21: Total abundance (no. ind. per m2) of dominant benthic macrofaunal taxonomic groups sampled at eight stations in St Helena Bay in 2001, 2007 and 2012. Data for sites SH31 and SH50 have been omitted from the 2001 adjusted graph to enable the data for the remaining sites to be plotted on the same scale as for the other sampling events (2007 and 2012).

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of in

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of in

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2

POLYCHAETA PENNATULACEA

MALACOSTRACA HOLOTHUROIDEA

GASTROPODA BIVALVIA

ACTINARIA

Page | 27

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.

0

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700

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2007

Tota

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s (g

) p

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m2

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POLYCHAETA

PENNATULACEA

MALACOSTRACA

GASTROPODA

BRACHYURA

BIVALVIA

ASCIDIACEA

ANTHOZOA

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2012

Tota

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s (g

) p

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POLYCHAETA

PENNATULACEA

MALACOSTRACA

GASTROPODA

BRACHYURA

BIVALVIA

ASCIDIACEA

ANTHOZOA

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2001

Tota

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) p

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POLYCHAETA

PENNATULACEA

MALACOSTRACA

GASTROPODA

BRACHYURA

BIVALVIA

ASCIDIACEA

ANTHOZOA

2001

2012

2007

Page | 28

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.

Figure 23. Average abundance per m2 of benthic macrofauna sampled in the bay and estuary in 2007 and

2012, indicated by functional groups.

Figure 24. Average abundance per m2 of benthic macrofauna sampled in the bay and estuary in 2007 and

2012, indicated by taxonomic groups.

0

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3000

2007 2012 2007 2012

Bay Estuary

Ave

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DETRITIVORE

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OCTOCORALLIA

NEMERTEA

MALACOSTRACA

GASTROPODA

CLITELLATA

BRACHYURA

BIVALVIA

ACTINARIA

Page | 29

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.

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

Group average

Old

_S

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4

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Ne

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SH

49

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3

Old

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H6

Old

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H2

1

Old

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0

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9

Old

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H3

0

Ne

w_

SH

50

Ne

w_

SH

30

Ne

w_

SH

31

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_S

H5

5

Ne

w_

SH

55

Ne

w_

SH

54

Ne

w_

SH

21

Ne

w_

SH

53

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_S

H2

6

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w_

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6

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2

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w_

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12

Samples

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80

60

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Sim

ila

rity

Transform: Fourth root

Resemblance: S17 Bray Curtis similarity

Year2007

2012



Year2007

2012

Similarity20

40

Old_SH6

Old_SH12Old_SH21

Old_SH26

Old_SH29

Old_SH30

Old_SH31

Old_SH46

Old_SH47Old_SH48

Old_SH49

Old_SH50

Old_SH53

Old_SH54

Old_SH55Old_SH56

Old_SH57

New_SH6

New_SH12

New_SH21

New_SH26

New_SH29

New_SH30New_SH31

New_SH46

New_SH47

New_SH48New_SH49

New_SH50

New_SH53

New_SH54New_SH55

New_SH56

2D Stress: 0.16

Page | 30

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.

Species Taxonomic group

Functional group

Change in abundance Contrib% Cum.%

Capitella capitata Detritivore Polychaeta Decreased (Absent in 2012)

20.9 20.9

Upogebia africana Filter Feeder Malacostraca Increased 9.86 30.76

Orbinia angrapequensis

Detritivore Polychaeta Increased (Absent in 2007)

8.77 39.53

Ceratonereis erythraeensis

Predator Polychaeta Decreased (Absent in 2012)

7.62 47.15

Spiroplax spiralis Detritivore Brachyura Increased (Absent in 2007)

5.3 52.45

Nereis spp. Detritivore Polychaeta Increased (Absent in 2007)

5.1 57.55

Callichirus kraussi Detritivore Malacostraca Increased 4.87 62.41

Glycera convoluta Predator Polychaeta Increased (Absent in 2007)

3.46 65.87

Grandidierella lutosa Detritivore Malacostraca Decreased (Absent in 2012)

3.38 69.25

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.

Page | 31

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.

Page | 32

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


SH12 1.9 not sampled 2.0




SH16 not sampled not sampled 1.6










SH46 not sampled 1.0 1.3 Increase

SH47 not sampled 0.7 1.3 Increase

SH48 not sampled 0.9 0.6 Decrease









Page | 33

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.

Page | 34

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).



TON

8E-2

0.32

0.56

0.8

SH4

SH6SH12

SH13

SH14SH15SH16 SH21

SH26 SH27

SH29

SH30

SH31

SH33

SH36

SH46

SH47

SH48

SH49

SH50

SH51

SH53

SH54

SH55

SH56

2D Stress: 0.18



Depth

7

28

49

70

SH4

SH6SH12

SH13

SH14SH15

SH16 SH21

SH26 SH27

SH29

SH30

SH31

SH33

SH36

SH46

SH47

SH48

SH49

SH50

SH51

SH53

SH54

SH55

SH56

2D Stress: 0.18



Mud

4

16

28

40

SH4

SH6SH12

SH13

SH14SH15

SH16 SH21

SH26 SH27

SH29

SH30

SH31

SH33

SH36

SH46

SH47

SH48

SH49

SH50

SH51

SH53

SH54

SH55

SH56

2D Stress: 0.18



TOC

0.9

3.6

6.3

9

SH4

SH6SH12

SH13

SH14SH15SH16 SH21

SH26 SH27

SH29

SH30

SH31

SH33

SH36

SH46

SH47

SH48

SH49

SH50

SH51

SH53

SH54

SH55

SH56

2D Stress: 0.18

Page | 35

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.

Page | 36

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).

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).

-30

-20

-10

0

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30

19

96

19

97

19

98

19

99

20

00

20

01

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20

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20

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10

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11

20

12

Sou

ther

n O

scill

atio

n In

dex

La Niña

El Niño

http://www.bom.gov.au/climate/glossary/soi.shtml

Page | 37

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.

Page | 38

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

Page | 39

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.

Page | 40

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.

Page | 41

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.

Page | 42

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

Page | 43

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

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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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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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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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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)

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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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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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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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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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St Helena Bay State of the Bay 2012 - Anchor Environmental