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KNYSNA ESTUARY: Ecology


  Flora
Phytoplankton/Diatoms
The phytoplankton biomass has not been investigated but the clarity of the water suggests that it is low.  A total of 39 species were identified by Korringa in phytoplankton samples taken for Thesen's Wharf, The Heads, Featherbed Bay and 2,5 miles offshore.

The phytoplankton has not been studied in detail but a number of species of diatoms have been recorded.  These include Skeletonema costatum, species of Chaetoceros, Coscinodiscus, Navicula and Pleurosigma.  Among the dinoflagellata Noctiluca miliaris is abundant at times and species of Ceratium and Peridinium are common.  (Ref 1)

Algae
Attached algal vegeation is rich.  The rocky banks at The Heads are colonised by an unusually wide variety of algal macrophytes.

The species of algae present include Gelidium pristoides, Ulva lactuca, Enteropmorpha sp. and Zonaria tournefortii.  Algae present within the lagoon include Ulva spp.  Enteromorpha spp. and Chaetomorpha spp.  The alga Zonaria tournefortii is common in Zostera beds.  Algae collected at The Heads, include Splachnidium rugsosum, Gelidium pristoides, Codium duthiae, Ulva rigida, Bryopsis sp.

The dry biomass of Ulva east of Thesen's Causeway was 61,5 g/m2.  While the ash-free dry biomass of Ulva east of Thesen's Causeway was 241 g/m2.  Patches of Ulva, Entermorpha and Chaetomorpha occur along the banks of tidal channels and on the surface of the mud on the lower shore.  The alga Bostrychia vaga forms tufts entangling the bases of intertidal saltmarsh plants. (Ref 1)

Aquatic Vegetation
From mid-tide down to the lowest level of spring-tide and even below, the dominant plant is the sea-grass Zostera capensis.  It grows all over the estuary up to the Old Drift.  It is most luxuriant at the low water of spring-tide level on shelving mud banks where the salinity is about 30 parts per thousand.  It is sometimes mixed with Halophila ovalis.  The mean dry biomass of Zostera is 67,5 g/m2 at Ashmead and 238,4 g/m2 north of Leisure Island.  The mean dry biomass of Zostera near the sewage outfall is 135 g/m2, the ash free dry biomass of Zoster in this area was 125 g/m2.  From Westford Bridge upstream Ruppia maritima becomes common eventually largely replacing Zostera at the Old Drift.  (Ref 1)
Semi-aquatic Vegetation
While Zostera beds predominate near the low-tide mark, borad saltmarshes comprised of various other species occur at higher levels.  Spartina maritima is abundant at mid levels and species of Sacocornia, Triglochin, Chenolea, Limonium and Plantage carpet the upper levels of the saltmarshes.  Near the high spring-tide mark, Juncus kraussii appers and it covers extensive areas of mud banks in the upper reaches of the estuary.  Biomass values for these macrophytes are high but primary production has not been measured.

The highest intertidal areas are carpeted with a dense growth of Limonium scabrum or Chenolea diffusa.  These species tend to form large patches covering areas of several hundred square metres each rather than a continuous band.  Limonium pathches extend down to 15 cm below high water of spring tide.  Below this it forms mixed communities with Chenolea, Sarcocornia and Triglochin but with Limonium still dominant in patches down to 20 cm high water of spring tide.  Slightly lower, occasional isolated plants of Limonium are each enmeshed in tufts of the alga Bostrychia vaga.  Sarcocornia decumbens tends to border the patches of Limonium but not spreading as widely in area. The densest patches are in the region of 15 -20cm below high water of spring tide.  Below this there are mixed patches of Chenolea and Limonium with Sarcoconia.  The lowest isolated specimens of Sacrcoconia were observed at 50 cm below high water of spring tide in the Triglochin zone.  Chenolea diffusa covers patches up to several hundred square metres in area between about 10-20 cm below high water of spring tide. Isolated plants merge into Triglochin zone about 25cm below high water of spring tide.  Chenolea appears to tend to favour the margins of channels and the slopes of humps perhaps favouring better drainage.  The lowest specimens occur in humps in the Triglochin zone at levels between 20-30cm below high water of spring tide.   (Ref 1)

Terrestrial vegetation
On the banks of the lagoon above high water of spring tide a large number of species of plants appear which are tolerant of marginal conditions including occasional salt spray.  Away from the immediate shore there is dense bush or scrub forest on the slopes.  The coastal bush on the Eastern Head (Foundation Point) and Western Head (Brenton Head) includes melkbos Sideroxylon inerme, camphor bush Tarconanthus camphoratus, taaibos Rhus glauca, assegaihout, Curtisia dentata, aloe Aloe ferox, num-num Carissa bispinosa, bietou Chrysanthomoides monilifera, blombos Metalasia muricata, polygala Poilygala sp., asparagus Asparagus sp. and buchu Agathosma sp.  (Ref 1)

Acock's map of the Veld types of South Africa indicates that the whole of the catchment and the surrounding of Knysna river and estuary fall in the Knysna Forest.  However, the crest of the hills stretching west from the western or Brenton Head support plant communities which appear to be representative of Coastal Fynbos although in Acock's map this is only represented west of Mossel Bay.  The vegetation of the Knysna Forest has been exhaustingly described by Phillips in this nmonograph on forest succession and ecology in the Knysna region.  Acocks points out that it is probable that the importance of Fynbos in the forest succession may be due to a large extent to carlesss exploitation.  He considers that sub-tropical grassveld might have been more important in the past in this region with high, well-distributed rainfall, sour sandy soils and vigorous growth. (Ref 1)

The composition of the forests is described by Von Breitenbach (1974).  There are about 125 tree and major shrub species of which about 100 are trees.  Throughout the Knysna area invasive alien vegetation has become a major problem.  Acacia saligna, Acacia cyclops, Acacia longifolia and Acacia melanoxylon have invaded large areas, sometimes eliminating the natural vegetation in the coastal belt.  In the Outeniqua Mountains Pinus pinaster and Hakea sericea now seriously threaten the survival of the rich Cape flora.   (Ref 1)
 

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Fauna
Zooplankton
The mouth of the Knysna Estuary is inhabited by neritic species while further up estuarine species dominate the zooplankton.  Among the copepod genera Paracalanus is abundant in the lower reaches while Pseudodiaptomus and Acartia species become abundant futher up.  Day has described the zooplankton as rather poor and suggested that plankton is not a major source of food in the estuary.  He pointed out that there are relatively few plankton feeders. 

The most common species of zooplankton recorded were Oithona nana, Paracalanus parvus, Harpacticus gracilis and Euterpina acutifrons, while Oithona oculata and Oncaea subtilis also appeared in sample from a cold current. (Ref 1)

The dominant copepods through most of the estuary are Acartiella natalensis, Paracartia longipatella, Paracalanus crassirostris, Paracalanus parvus, Oithona nana, Pseudodiaptomus nudus and Harpacticus gracilis.  These species occur in most estuaries on the south and east coasts of South Africa.  In addition various neritic marine forms such as Clausoclanus furcatus, Centropages brachiatus, Pleuromamma abdominalis, Euterpina acutifrons and various species of Corycaues penetrate some distance into the estuary.  The Knysna estuary is unusual among South African estuaries in being a deep open estuary in which clear blue open water penetrates on each high-tide.  The blue water during spring-tides penetrates to well beyong the railway bridge mixing with the more turbid green water in the shallows.  This facilitates the penetration of neritic marine species and limits the survival of purely estuarine species such as Pseudodiaptomus hessei and varous Hapacticoids.  These latter species are abundant in the upper reaches from the Belvedere area right up to Charlesford Rapids.  The true estuarine plankton species can only survive in areas where the residence time of the water is sufficiently long for them to be able to complete their life cycles before being swept out to sea by the tides.

At times, very cold upwelled water apppears at Knysna and penetrates into the lagoon.  At such times various deep ocean species appear such as Oncaea subtilis, Oithona oculata and Microsetella rosea.  These are oceanic pelagic species which do not usually form part of the estuarine or neritic marine plankton community. (Ref 1)

The Chaetognatha are represented by a species of Sagitta.  Large numbers of larvae of benthic invertebrates appear in the plankton particularly during the spring and summer months.  These include polychaete larvae, larvae and post larval stages of gastropods and lamellibranchs, ophioplutues and other echinoderm larvae, copepod nauplii, cirripede nauplii, cirripede cyprids, crab zoea and megalopa larvae and mysis larvae of decapods.  Several species of Ostracoda have been recorded.  Among the Mysidacea Gastrosaccus brevifissurs and Leptomysis tattersali occur in the lower estuary while Mesopodopsis africana and Rhophalopthalamus egregius occur in the upper estuary.  Cumacea include Iphinoe truncata. Amphipoda include Paramoera capensis, Lysianassa ceratina and Corophium triangonyx.  Isopoda include,Eurydice longicornis, Janira capensis, Paridotea ungulata and Exosphaeroma spp.  Various fish eggs and larvae appear and occasionally small fish such as the pipefish Syngnathus acus. (Ref 1)
As might be expected in a clear open estuary penetrated by ocean water at every tide there are more neritic marine species of phytoplankton and zooplankton that in most South African estuaries.  Organic detritus particles are abundant in most plankton samples sometimes even exceeding the plankton in abundance.  It is clear that the detritus derived from the attached plants of the saltmarshes plays an important role in this estuary.  Nanoplankton is particularly abundant in the upper reaches of the estuary and it appears to be the major food source of the dominant copepod in that area Pseudodiaptomus hessei(Ref 1)

Results of numerous studies indicate that total phytoplankton biomass within the estuary is low, generally < 2 mg chl-a m-3, which can be linked to the low macronutrient availability within the system. Statistical analyses reveal no significant spatial differences in the total phytoplankton biomass along the length of the estuary (Grindley, 1985; Allanson et al., 2000a). Wind generated coastal upwelling in the marine environment appears to play an important role in determining the phytoplankton biomass within the Knysna Estuary. Cold, nutrient rich water penetratong into the estuary is associated with a dramatic increase in the total phytoplankton biomass within the estuary with levels of up to 18.4 mg chl-a m-3, being recorded (Allanson et al., 2000). The freshwater inflow from the Knysna River is also associated with increases in the phytoplankton biomass as riverine inflow represents the most importance source of macronutrients necessary to sustain the growth of the phytoplankton. These results highlight the importance of catchment management strategies in maintaining the health of the Knysna Estuary.  A comprehensive description of the zooplankton community within the Knysna Estuary is given byGrindley (1985). The zooplankton community structure within the Knysna Estuary demonstrates strong horizontal patterns reflecting the hydrodynamics of the system. Neritic (oceanic) species dominate in the lower and middle reaches of the estuary while true estuarine species predominate only in the upper reaches (Grindley, 1985). The zooplankton community throughout the estuary is numerically dominated by copepods. Among the copepods, species of the genera Oithona, Paracalanus, Oncaece and Pseudodiaptomus dominate in the lower and middle reaches while in the upper reaches the estuarine copepods on the genus Psuedodiaptomus and Acartia are most numerous. In addition to the copepods, several additional neritic species of chaetognath, mysid and gelatinous zooplankton have also been recorded within the system.  (Ref 5)

Aquatic Invertebrates

The benthic macrofauna includes 310 species with rich beds of Upogebia africana and numbers of Arenicola loveni, Solen corneus, Atrina squamifer and other bivalves.  The gastropods Asiminea globulus, Hydrobia sp. and Nassarius kraussianus are abundant.  There are numerous amphipods and isopods and the saltmarsh vegetation harbours Sesarme catenta, Cleistostama spp. and other crabs.

Corresponding to physical changes along the length of the estuary there are a series of faunsitic divisions.  Day et al. (1952) pointed out that salinity changes, current strength, pH and temperature may all be important, but that the environment must be considered as a whole.  Any one factor only becomes important when the limits of tolerance for that factor are reached.  Four basic divisions of the Knysna Estuary were distinguished:
  1. Knysna Heads
  2. the Lagoon from Leisure Island to beyond The Point
  3. the Westford channels to the Old Drift and 
  4. Charlesford Rapids.

It was shown that the diversity of species declines progressively as one passes upstream from The Heads.  While the number of species present at Thesen's Island is roughly similar to that at The Heads, the component species are notably different.  Thus over half the species at The Heads do not extend to The Heads.  A characteristic lagoon fauna was recorded from Leisure Island to The Point.  The percentage of typical seashore species decrease progressively up the lagoon and none reaches Charlesford Rapids.  (Ref 1)

The Pansy Shell, Echinodiscus bisperforatus occurs and bredds on particular sand banks in the main channel of the lagoon.  They are popular with visitors but their numbers appear to have declined alarmingly in recent years, dejpsite conservation legislation.

Knysna is a major supplier of oysters.  Oyster larvae settling on collectors placed inside the Knysna Lagoon come from parent stocks in the sea.  They are carried by the counter-current flowing very close inshore, and are washed in to the lagoon with the flood tide.  Therefore settling takes place close to the entrance of the lagoon and at about high-tide.  Conditions in the central part of the lagoon, above the Rail Bridge, are not suitable for oyster larvae, but spat and older oysters may thrive there because of the high food content of the water.

Three species of indigenous oysters appear to occur at Knysna.  Of these the common South African oyster, Crassostrea margaritacea, an oviparous species, is considered the most promising of oyster cultivation.  The small  Ostrea algoensis should be considered as a  'weed-oyster' because of its limited dimensions and slow growth.  By means of proper timing of the placing of collectors it is possible to avoid this species settling in appreciable numbers.  The red oyster, Ostrea atherstonei, another larvipqarous species, is very interesting because of its rapid growth and perfect shape.  Previously this species was only known from fossil deposits.  It occurs at many places along the coast of South Africa, including the west coast, but the numbers settling at Knysna are not of commercial significance, so that it will be extremely difficult to build up a stock of this oyster.  In addition to the indigenous oysters, the Pacific oyster Crassostrea gigas was introduced for cultivation in 1973. (Ref 1)

The Pacific oyster C.gigas was introduced in 1973.  Present production is two million per annum from Knysna Lagoon. This represents approximately 200 tonne or more per hactare per year.  Cultured seed oysters are procuded at the Fisheries Development Corporation hactchery and supplied to the oyster farming industry.  An intertidal tray system is currently employed for further growth until harvesting.  The Portuguese oyster, Crassostrea angulata, grows and fattens satisfactorily in the Knysna Lagoon.  (Ref 1)

The Knysna Estuary is home to a number of critically endangered invertebrate species including the Knysna seahorse (Hippocampus capensis), the pulmonate impet, Siphonaria compressa, and the Pansy shell (Echinodiscus bisperforatus) (Grindley, 1985; Angel et al., 2006). As a consequence, the Knysna Estuary ranks very highly in terms of its conservation importance and the preservation of its fauna and flora would ensure that nearly 43% of South Africa estuarine biodiversity would be conserved (Turpie, 2000). (Ref 5)

Insects
Insects have not received detailed study but those collected included chironomid midges and their larvae, dragonflies, mayflies, kelp flies, staphylinid beetles and other beetles, house flies and water boatmen. (Ref 1)
Fish
More than 200 species of fish are reputed to be found in the lagoon which is, inter alia, the home of a rare seahorse, Hippocampus capensis.

During the University of Cape Town ecological studies, over 50 species of fish were identified.  Most fo these were juveniles of marine species but anglers also take large kob, leervis, spotted grunter, white steenbras, elf and even stenohaline marine fish such as Sparodon durbanensis on occasion.  Forage fish such as Gilchristella, Hepsetia breviceps and three or four species of mullet are plentiful.  (Ref 1)

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The white stumpnose is one of the commonest fishes and feeds on mussels and small crustaceans particularly such as amphipods.  The white steenbras grubs in the sand for polychaetes such as Orbinia and Nephthys spp. and small crustaceans particularly Pontogeloides and Urothoe.  Large specimens develop teh cpaacity of blowing holes in the sand to feed on Upogebia and Arenicola.  The kabeljou appear when they are about 15cm long.  The feed ona variety of small crustceans including mysids, shrimp, Upogebia and Hymenosoma by they do not seem fo feed on the infauna of muddy bottoms.  Large specimens predatory and feed on Sepia and Mugil spp.  Adult kob extend the whole length of the estuary including the upper reaches.  The leervis Lichia amia is a vicious predator even when small, feeding on shoals of post-larval mullet in the shallows.  They extend into the upper reaches of the estuary and as adults feed on Mugil spp. Hyporhamphys and juvenile Lithognathus and Rhabdosargus.  They are swift predators and form the final link in the food web illustrated in a diagram of the trophic relations within Knysna Estuary.  (Ref 1)
Because of the permanently open wide rocky entrance, the area near The Heads includes typical marine fishes.  The width of the mouth is such that there is no barrier to shoals of pilchards, anchovies, etc. and their attendant predators, so they enter freely.  As a result there have been many records of fishes unusual for estuaries.  At times, the upwellign on the south coast drives numbers of species into the lagoon. Some of these are thrown ashore numbed or dead.    (Ref 1)
Amphibians and Reptiles
Snakes and lizards that have been observed around the estuary include the russet snake, the night adder, the puff adder, a skink, and a gecko.  There are also a number of toads and frogs that occur in the Knysna area.   (Ref 1)
Birds
A great diversity of birds frequent the Knysna area and the Knysna Estuary and its saltmarshes in particular.  The Knysna loerie, Tauro corythaix,  is well known.(Ref 1)
Mammals
The largest mammal found in the Knysna area is the African elephant Loxodonta africana africana. Mammals that are seen from time to time around the estuary include the Cape dassie, the Cape Grysbok, Cape Grey mongoose, the Cape dune mole and the Cape fruit bat. (Ref 1)

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Ian Flemming Photography 

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