Management Plan for the GARDEN ROUTE NATIONAL PARK

APPENDICES

APPENDIX A

PRESIDENTS MINUTE OF 17 March 2010

management plan GRNP Appendices 1 management plan GRNP Appendices 2

APPENDIX B

MARINE MANAGEMENT PLAN

Undergoing completion, to be inserted

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APPENDIX C

VEGETATION OF THE GRNP

FRESHWATER & ESTAURINE

Phytoplankton

Phytoplankton populations in the Wilderness lake systems are low (Robarts 1973), as confirmed by chlorophyll-a determinations undertaken in Swartvlei Lake in 1976 (Coetzee 1978) and 1977- 78 (Howard-Williams & Allanson 1981a). Three major categories of phytoplankton have been recorded in Swartvlei Lake viz. diatoms, flagellates and dinoflagellates (Robarts 1976) with the diatom Coscinodiscus lineatus the most abundant species. Other common diatom species include Chaetoceras wighamii, Cocconus scutellum, Grammatophora oceanica, Grammatophora sepentia, Navicula pseudony, Raphoneis mirabunda, Raphoneis superba and Synedra tabulata (Robarts 1973). Flagellates and dinoflagellates generally form a relatively minor part of the phytoplankton biota (Robarts 1973), though short-lived blooms do occasionally occur.

Thirty nine phytoplankton species have been identified by Korringa (1956) in Knysna Estuary. Phytoplankton biomass has never been investigated, though Day (1981) maintains that the clarity of the water in Knysna suggests that it is low. Grindley (1985) lists some of the diatoms recorded in Knysna Estuary, and Grindley (1976) and Grindley & Eagle (1978) list abundant dinoflagellata.

Algae

Principal genera of epiphytic algae occurring in the Wilderness lakes include Enteromorpha, Lyngbya, Cladophera, Percursaria, Cocconeis, Ectocarpus, Polysiphonia, Chondria and Hypnea (Howard-Williams 1980; Howard-Williams & Liptrot 1980). The rocky banks at the mouth of Knysna Estuary are colonised by a wide variety of attached algal macrophytes (Day 1981). Common species within the estuary include Gelidium pristoides, Ulva lactuca, Enteromorpha spp. Chaetomorpha spp. and Zonaria tournefortii (Day et al . 1952, Day 1967, Day 1981, Grindley 1976, Grindley & Eagle 1978, Grindley & Snow 1983).

Submerged aquatic plants

Submerged macrophytes are widespread in the Wilderness lake systems and consist predominantly of pure and mixed stands of Potamogeton pectinatus , Charophyta and filamentous algae (Howard- Williams & Liptrot 1980, Weisser & Howard-Williams 1982, Whitfield et al. 1983) with Ceratophylum demersum (hornwort) having become widespread and abundant in several lakes during the 1990’s. In the estuaries Ruppia spirralis and Zostera capensis predominate. Macrophytes do not generally occur in waters deeper than 3.0m (Whitfield 1984). The production

management plan GRNP Appendices 4 of organic matter by submerged aquatic plants was calculated in 1978-1979 as 1.84 x 10 6 kg y -1 (dry weight) (74% of total production) in the Swartvlei system (Howard-Williams & Allanson 1979), and 334 100 kg y -1 (17% of total production) in the Touw system (Howard-Williams 1980). The bulk of this organic matter is produced in the shallow periphery of the waterbodies, in waters to a depth of three meters.

Both long- and short-term changes occur in the abundance, biomass and distribution of submerged aquatic plants, with substantial declines in the abundance and distribution of species were recorded in most waterbodies between 1973 and 1982 (Davies 1982; Weisser & Howard-Williams 1982; Whitfield 1982; Weisser et al . 1992), though in 1991 the standing crop of submerged macrophytes in most waterbodies was found to be similar or greater than those recorded prior to 1973 (Russell 1996a). Only in Rondevlei has the biomass of submerged aquatic plants increased consistently with time (cf. Howard-Williams 1980; Hall 1985a, 1985b; Russell 1996a). Weisser et al . (1992) suggest that aquatic vegetation proceeds through cycles, from an aquatic macrophyte dominated system through a phytoplankton dominated system and back to a macrophyte dominated system. Theorised reasons for periodic declines in aquatic plants include fungal diseases (Howard- Williams 1980), changing nutrient status of the waterbody (Hall 1985a, 1985b; Weisser 1979), shading by Enteromorpha (Hall 1985a) and dinoflagellate blooms (Coetzee & Palmer 1982), reduced water transparency resulting from the influx of turbid water (Whitfield 1982; Allanson & Howard-Williams 1984), and reduced calcium: magnesium ratios in the water column as a result of persistent flooding (Allanson & Howard-Williams 1984).

Howard-Williams & Allanson (1979) described how a 10% decline in submerged aquatic plants in Swartvlei Lake would result in a 20% decline in the total food production. The ratio of food production to consumption in Swartvlei Lake is approximately 1:1 (Howard-Williams & Allanson 1979), while in the estuary utilisation of organic material by the biological community is greater than the production rate, a deficit made larger by the export of Zostera leaves during the tidal phase (up to 500 kg dry mass per tide). Hence any loss in primary production would result in a corresponding reduction in the number of consumer organisms (see section 3.4.1 Suspensoids).

The role played by aquatic plants in cycling nutrients between the water column and the bottom sediments has been extensively studied in the Swartvlei system (see Howard-Williams & Allanson 1979). In Swartvlei Lake production rate by submerged aquatic plants is high (e.g. P. pectinatus = 16 g m -2 day -1) (Howard-Williams 1978). In this waterbody no evidence of phosphorous limitation was found in Potamogeton tissues (Howard-Williams 1977), whereas the large algae, Chara and Cladophera spp. were found to be nutrient limited. Potamogeton does not act as an efficient “nutrient pump” (Howard-Williams & Allanson 1979), thus the release of nutrients from the rooted aquatic plants is through decomposition which takes place rapidly predominantly in spring and early summer (Howard-Williams & Davies 1979). Bacteria as opposed to fungi are primary colonisers on leaves resulting in decomposition (Howard-Williams et al . 1978; Robb et al. 1979). It is unlikely that P once cycling in the macrophyte community, would become transferred to the circulation in the open lake (Howard-Williams & Allanson 1981b). The rate of uptake of nutrients from the water column by epiphytic algae is approximately ten orders of magnitude greater than

management plan GRNP Appendices 5 that of the Potamogeton (Howard-Williams 1977, 1981), thus enrichment of the water with P and N compounds would be expected to result in the growth of epiphytic algae. The sediments of Swartvlei Lake also act as a major sink to plant nutrients, absorbing up to 60 % of all phosphorous inputs into the system (Howard-Williams 1977, 1981). In a recent reworking of Swartvlei aquatic macrophyte data collected in the 1970’s a model was developed to simulate the decomposition process, and the P and N content of the remaining biomass (Asaeda et al . 2000). Model outputs were comparable with measurements of dry matter and nutrient loss as observed by Howard- Williams & Davies (1979). The model can be used to simulate reductions in nutrients in shallow lakes following the harvesting of Potamogeton .

In Swartvlei Estuary Z. capensis acts as a nutrient pump during the day adsorbing nutrients from the sediments and secreting them into the surrounding water (Howard-Williams & Allanson 1979). It is postulated that there are two pathways of the take-up of nutrients, the first being a sediment- water exchange system of plant nutrients, the second being that the large mats of Enteromorpha algae take up P during the day faster than the rate at which it is released by the Zostera , and then in turn release P compounds during the night. During the tidal phases there is a net import of P into the system, primarily in the form of particulate matter from the sea (Howard-Williams & Allanson 1979). There is no evidence of a long-term accumulation of phosphate in Swartvlei Estuary (Howard-Williams & Allanson 1979) which indicates that accumulated nutrients must be periodically removed. The strong currents which follow the opening of the estuary remove large quantities (1.6 tonnes over 20 tidal cycles / 18% of Zostera bed biomass per month) of aquatic plants (primarily Zostera and Enteromorpha ), detritus and sediments (Whitfield 1988b), which constitute a loss of plant nutrients from the system. In this manner a long-term equilibrium is maintained. Weed drift also helps redistribute nutrients within Swartvlei Estuary, with the resulting detritus layer supporting large numbers of detritus feeders.

In the Knysna Estuary the dominant plant on shelving mudbanks below the mid-tide level is the eel-grass Z. capensis (Grindley 1985). The mean dry biomass of Z. capensis is 67.5 g m -2 at Ashmead, and 238.4 g m -2 north of Leisure Island (Grindley 1976). In places Halophila ovalis grows in association with Z. capensis . In the upper reaches of the estuary, from Westford Bridge upstream, Rupia maritima becomes common, eventually largely replacing Z. capensis at the Old Drift (Grindley 1985).

The Groot River (West) estuary is the largest and probably the only estuary in the Tsitsikamma region that has notable stands of submerged aquatic plants, with stands of Ruppia maritima and R. spiralis occurring in the creek west of “The Island” (Morant and Bickerton 1983).

Emergent aquatic plants

Occurrence and distribution of emergent aquatic plants are given in Jacot-Guillarmod (1979, 1981, 1982), Weisser & Howard-Williams (1982), Whitfield et al . (1983) and Russell (2003). Phragmites australis has been calculated to produce up to 303 000 kg y -1 organic matter (dry weight) (12% of total production) in the Swartvlei system (Howard-Williams & Allanson 1979),

management plan GRNP Appendices 6 and 1.08 x 10 6 kg y -1 (55% of total production) in the Touw system (Howard-Williams 1980). Organic matter production by other semi-aquatic plant species is substantially lower, with Schoenoplectus scirpoideus producing 351 600 kg y -1 (18.0% of total production), and Typha capensis 182 800 kg y -1 (9.4% of total production) in the Touw system (Howard-Williams 1980). No estimates of annual biomass by S. scirpoideus and T. capensis are available for the Swartvlei system.

In the Touw system Howard-Williams (1980) found that P. australis and S. scirpoideus decreased in abundance and biomass from the Touw River Estuary/Serpentine towards Rondevlei, whereas T. capensis increased in abundance and biomass in the same direction. Mapping of the distribution of plants in the 1970s (Weisser & Howard-Williams 1982) and 1990s (Russell 2003) indicated localised increases in P. australis, T. capensis, Scrub or trees, and Grass or Fields, and decreases in Juncus kraussii, Schoenoplectus scirpoideus and Low scrub or fynbos in the Touw system. Probable causes of change in the distribution of wetland plants include the natural tendency of plants to colonise new areas, as well as anthropogenic manipulation of physical, chemical and biological processes, including the cessation of disturbance by large herbivores, water-level stabilisation, changes in soil salinity and the accumulation of plant litter within wetland areas (Russell 2003).

The intertidal wetlands of the Knysna Estuary cover an area of 1000 ha (Maree 2000) extending landward of the mid-tide level. Fifty four plants species have been collected in Knysna saltmarshes, of which 27 occur exclusively in this habitat (Maree 2000). Spartina maritima is abundant at the mid-tide level, whereas Sarcocornia, Triglocin, Chenolea, Limonium and Plantago species predominate in the upper levels of the saltmarshes (Grindley 1985). Juncus kraussii occurs near the high spring-tide level, and covers extensive areas of mudbanks in the upper reaches of the estuary (Grindley 1976). Maree (2000) has mapped the distribution of the main types of plant communities as occurring in 1990.

Estimates of biomass production (g m -2 dry mass) by saltmarsh plant species in communities on the eastern side of the estuary are given by Grindley (1976), Grindley & Eagle (1978) and Grindley & Snow (1983). The role of saltmarsh vegetation as a source of detritus has been discussed by Grindley (1976). Maree (2000) identified several threats to saltmarsh communities, including trampling by people, vehicles and moored boats, bait collecting, constructions, high velocity inflows from drains, and high silt loads.

In Groot River (West) estuary, the largest estuary in the Tsitsikamma region, wetland species such as Mariscus thunbergii , Juncus kraussi and Phragmites australis are found on the western bank, while Samolus porosus is abundant on the island and eastern bank (Morant and Bickerton 1983). Based on aquatic and semi-aquatic vegetation, this estuary had an intermediate botanical importance rating compared to 31 other estuaries in the Cape Province, while that of the Salt River estuary was low (Coetzee et al . 1997). The vegetation of the freshwater sections of these and other river systems in the park is poorly studied. However, stands of Prionium serratum (palmiet) have been noted in the Salt- (De Moor et al. 2004), Storms- and Palmiet rivers.

Truly emergent aquatic plants are a rarity in the Fynbos and Forest biomes. Common, however, is Aponogeton distachyos that is commonly found in slow running rivers or river pools in the forest.

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Another Aponogeton cf. species, as well as a diminutive Cyperaceae where found in a vlei in Petrus Brand Nature Reserve. Marsilea schelpeana, Nymphoides species and Nymphaea nouchali occur in Goldblatt & Manning (2000) and the GRNP species list. See Species List for GRNP for complete list of emergent plants. Some common species that are marginally aquatic, and are closely related to water or water courses are Prionium serratum , Isolepis prolifera and Cotula coronopifolia.

Marsilea schelpeana and Prionium serratum occurs on the Red Data List for plants

INDIGENOUS FORESTS

Mucina and Rutherford (2006) follow the latest Department of Water Affairs and Forestry (DWAF) national forest type classification system for South African indigenous forests (Von Maltitz et al. , 2003) and identify two national forest types occurring in the Garden Route, viz. Southern Cape Afrotemperate Forest and Western Cape Milkwood Forest.

Southern Cape Afrotemperate Forest

The Southern Cape Afrotemperate Forests is the largest forest complex in southern Africa (approximately 60 500ha; Geldenhuys, 1991). They occur from sea level to altitudes of 1 000 metres or more, but most of the high forest occurs on the coastal plateau and foothills of the mountains. They display great variation in forest structure, species composition and species density due to the variations in topography, soils, microclimate and other factors. The forests occur in three distinct zones with distinct stand structure and species composition (Geldenhuys 1982). The mountain forests (mostly wet forests) are typical Afromontane forest. Dry forests or scrub forests occur on the coastal scarp or on the steep slopes of the river valleys (Phillips 1931). These forests are species-rich and include many species of Indian Ocean Coastal Belt forests (Moll & White, 1978 in Geldenhuys, 1982). The plateau forests are typically moist and medium-moist forests and include mainly species of Afromontane affinity.

A total of 465 plant species are known to occur in these forests. This is relatively high for the southern latitude of these forests and can be attributed to the range of diverse habitat types and dispersal corridors that linked these forests to the east over a very long period (Geldenhuys 1992a). There are several species that drop out along different zones from east to west within the southern Cape, and many species have a disjunct distribution within the area (Geldenhuys 1992b).

Following the pattern of the climatic zones, from the hot and dry zone of the littoral over the temperate zone to the cold and wet zone of the mountain ranges, Von Breitenbach (1968a-c, 1974) described eight climax forest types. Of these seven are significant in the management system. The classification is done according to the combined assessment of moisture regime, ground shrub and tree flora as well as features such as tree form and canopy height. However, sites are ecologically complex and produced by varying combinations of interchangeable factors. Forest types are therefore

management plan GRNP Appendices 8 crude, and of necessity artificial groups of areas which have similar site features. This forest type classification however effectively serves the purpose of providing area-specific information on resource nature and utilisation potential and constraints (e.g. ecological sensitivity) and as such strongly influences management objectives and practices.

The forest types, illustrated in Figure 1 below, are as follows: Very very dry Scrub (vvd-S) Found on sites with a very hot and dry climate, and where soil formation occurs in patches only and remains at an initial stage over a sandy, rocky or stony substratum. The scrub is about 2-5 m high and consists of smaller or larger groups of shrubs and occasional, stunted and gnarled, bushy trees, with low ground vegetation. The shrub flora is extremely rich in species. Among the most common shrubs that occur are Allophylus decipiens, Buddleia saligna, Cassine peragua, Chrysanthemoides monilifera and Osyris compressa . The shrubs are often intertwined, or overgrown by scramblers, such as Rhoicissus tomentosa and Scutia myrtina . The ground between the scrub groups is covered with dwarf-shrubs and herbaceous plants among which species of Agathosma and Cliffortia are prominent.

Very dry Scrub Forest (vd-SF) Occurs on sites with a fairly hot and dry climate, and on very shallow sandy soils, immature clay soils or loam soils that dry out rapidly. Structurally this type is characterised by a dense mixture of shrubs (3 to 6 m tall) and stunted, bushy trees (6 to 12 m tall). Large trees are occasionally found. Abundantly represented as small trees are Robsonodendron aethiopica and C. peragua . Other small trees include Apodytes dimidiata , Pterocelastrus tricuspidatus and Vepris undulata . Thorny species such as Gymnosporia heterophylla , Carissa bispinosa and Scutia myrtina abound. The forest floor is usually covered with dry leaf litter and is practically devoid of a herbaceous ground layer. Only where the otherwise dense canopy is somewhat open, and in marginal areas, groups of low ferns and herbs are found, such a Cheilanthes hirta, Pellaea calomelanos and Streptocarpus rexii . A particularly characteristic herb is Gerbera cordata .

Dry High Forest (d-HF) Found on sites with a moderately hot, dry climate and where soils are deep well-drained sands, or deep clay soils, or shallow loam soils over clay or stone, which dry out after summer rains. The forest is relatively dense and has an irregular canopy 10 to 18 m high. Among the trees forming the main canopy, Pterocelastrus tricuspidatus is usually the most frequent. Other common species are Apodytes dimidiata , Psydrax obovata, Podocarpus falcatus and P. latifolius . Tree regeneration is poor, but there is an abundance and variety of ground flora species constituted by ferns, grasses and herbs such as Dietes iridioides, Oplismenus hirtellus and large groups of Rumohra adiantiformis . The undergrowth is inundated by thorn species such as Carissa bispinosa and Scutia myrtina . Trichocladus crinitus appears in scattered groups.

Medium-moist High Forest (mm-HF)

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Occurs on sites with a temperate and fairly humid climate, and with shallow to moderately deep, dense loam soil over an impenetrable clay substratum: soils that, due to their impeded drainage, are fairly moist throughout the year. This type of forest has two relatively dense canopy layers above a very dense undergrowth layer and a generally scanty ground flora. The main canopy lies 16 to 22 m above the forest floor and is made up of the crowns of the large, dominant trees. The lower canopy layer lies about 10 m below the main canopy and consists of pole-sized trees of species found in the upper layer together with various other smaller species. The undergrowth reaches a height of 6 m. Olea capensis subsp. macrocarpa is the most common dominant; others include Podocarpus latifolius , P. falcatus , A podytes dimidiata , Olinia ventosa , Pterocelastrus tricuspidatus and Rapanea melanophloeos . In the lower canopy layer Gonioma kamassi is the most common. The undergrowth consists almost entirely of dense Trichocladus crinitus . Fine lianes are frequent in the lower storeys. Cynanchum ellipticum, C. obtusifolium and Zehneria scabra often bind the undershrubs together. Ground flora of small ferns, low grasses and herbs is very poor under the dense undershrub, but it becomes richer where the forest is open. Besides scattered patches of Oplismenus hirtellus , there is also abundant Dietes iridioides . Tree regeneration is prolific even under dense onderbos.

Moist High Forest (m-HF) Occurs on sites with a temperate and humid climate, and with moderately deep to deep loam soils over an impenetrable clay substratum. Soils, due to the low rate of evaporation, its density and the bad drainage or even waterlogging of its deeper layers, are moist throughout the year. There are three highly irregular, relatively dense crown layers, lying at 20 to 30 m, 12 to 20 m and 6 to 12 m above the forest floor. The middle canopy consists of immature upper canopy species together with the larger crows of a number of intermediate storey species. The lower storey consists of the sparse crowns of scattered understorey species. The dominants of the upper layer are mostly Olea capensis subsp. macrocarpa and Podocarpus falcatus , the latter sometimes attaining heights of just over 40 m. Others in the upper layer include Apodytes dimidiata , Ocotea bullata , Podocarpus latifolius , Rapanea melanophloeos and Nuxia floribunda . Pterocelastrus tricuspidatus , Curtisia dentata , Psydrax obovata and Platylophus trifoliatus occur in the middle canopy. Gonioma kamassi is usually plentiful, while among others Halleria lucida and Ochna arborea are quite frequent in the lower storey. In the undergrowth Trichocladus crinitus may occur in dense groups, but is generally scattered and open. Ground flora is abundant and dense communities of Plectranthus fruticosa, Piper capense and Rumohra adiantiformis often replace the undershrub in more open parts. Tree regeneration, however, is generally poor.

Wet High Forest (w-HF) Found on sites with a cool and wet climate, and on shallow to moderately deep sandy loam soils which are badly drained or even waterlogged under the thin upper layer and thus wet throughout the year. The forest consists of a medium-dense, generally two-storied stocking of narrow-crowned and slender-stemmed trees over a dense tree fern storey or luxuriant, shrub-interspersed ground vegetation. The general canopy is about 12 to 20 m high and irregular. A lower canopy may be determined about 6 m below the main layer and is made up of scattered small trees. Ocotea bullata , Cunonia capensis and Podocarpus latifolius are the dominant species. The lower canopy includes

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Curtisia dentata , Halleria lucida and Olea capensis subsp. capensis . A characteristic of wet forest is the presence of Cyathea capensis which may reach a height of 6 m. Tree regeneration is almost non- existent where ferns are plentiful, but profuse in more open areas. Trichocladus crinitus is sparse or often totally absent.

Very wet Scrub Forest (vw-SF) Found on sites with a very cool and wet climate, and with very shallow, peaty loam soils. The general canopy is 6 to 10 m high. About 3 m below the canopy is a shrub layer. Cunonia capensis is the dominant species, sometimes rising above the general canopy. Other trees include Ocotea bullata , Platylophus trifoliatus , Podocarpus latifolius , Rapanea melanophloeos and Virgilia oroboides . The shrub layer includes species such as Cyathea capensis , Diospyros glabra and Sparmannia africana . The ground flora consists of dense fern growth in which Blechnum sylvaticum, B. giganteum, B. australe, Gleichenia polypodioides and Todea barbara are prominent.

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Figure 1: Indigenous forest types of the Southern Cape and Tsitsikamma (Von Breitenbach, 1974).

Western Cape Milkwood Forest

Western Cape milkwood forest occurs as small patches at altitudes between 20 and 340m along the southern coast between Nature’s Valley and the Cape Peninsula (Mucina & Rutherford, 2006; Von Maltitz et al. , 2003). In the Garden Route, patches occur inter alia at Nature’s Valley, Keurbooms, Knysna, Goukamma and Groenvlei Forests in the Goukamma Nature Reserve, Sedgefield and Wilderness National Park.

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The forests are generally low, species-poor and often dominated by only one or a few canopy tree species with large stems and spreading crowns, usually Sideroxylon inerme and Celtis Africana or Apodytes dimidiata . The understorey may be open, but is often a shrub layer with a diverse range of species.

FYNBOS

Most of the Fynbos biome, and also the fynbos in the park, forms part of the Cape Floristic, Kingdom, the smallest of the six Floral Kingdoms of the world. It is categorized by a high plant species richness and endemicity (Low & Rebelo 1996). About 65% of the park is covered by natural fynbos, representing eight national types, comprising of three types of Sandstone Fynbos, two types of Sand Fynbos, and one type each of Shale Fynbos, Fynbos Shale Band Vegetation and Granite Fynbos, as described by Mucina & Rutherford (2006). The GRNP contains some 47% of the vulnerable South Outeniqua Sandstone Fynbos, some 3% of the endangered Knysna Sand Fynbos, some 16% of the Southern Cape Dune Fynbos, and some 4% of the endangered Garden Route Shale Fynbos (Mucina and Rutherford 2005).

Sandstone Fynbos

Together with Southern Afrotemperate Forest, Sandstone Fynbos covers the largest terrestrial area of the park), comprising of a range of Dry, Mesic and Wet Mountain Fynbos types ( vide Campbell 1985). It has been classified as South Outeniqua Sandstone Fynbos and Tsitsikamma Sandstone Fynbos on the southern slopes of the Outeniqua and Tsitsikamma Mountains respectively, with smaller areas of North Outeniqua Sandstone Fynbos (Mucina & Rutherford 2006). In the mountainous areas of both these types, pockets of Afrotemperate Forest are found in gullies and bergwind shadow areas where they are protected against fynbos fires (Geldenhuys 1991, 1994).

South Outeniqua Sandstone Fynbos is found on gentle to steep, mainly south-facing slopes in the Outeniqua Mountains, on acidic lithosol soils derived from Ordovician sandstones of the Table Mountain Group. Pierce (2003) further divide the type into Outeniqua Mountain Fynbos to the north and Outeniqua Plateau Fynbos, largely on the southern boundary of the park. It consists of a tall, open to medium dense shrub layer with a medium tall shrub understorey, and comprises of mainly proteoid and restioid fynbos and extensive ericaceous fynbos on the upper slopes (Mucina & Rutherford 2006). The type is also well represented as fynbos islands in the Afrotemperate forest in the park, originating from the expansion of forest vegetation during, isolating it from the mainland fynbos ( vide Bond et al . 1988, Midgley & Bond 1990). Floristic communities that form part of South Outeniqua Sandstone Fynbos have been described by inter alia Seydack (1977), Bond (1978, 1981), West (1989) and Vermeulen (1995). Important species include Widdringtonia nodiflora (small tree), Laurophyllus capensis , Leucadendron eucalyptifolium (tall shrubs), Berzelia intermedia , Erica densifolia (low shrubs), Carpacoce spermacocea , Centella affinis (herbs), and the graminoids Cannomois virgata and Restio triticeus . A large number of Erica species are

management plan GRNP Appendices 13 endemic to this type, including Erica unicolor , E. aneimena and E. gillii . Other endemic taxa include the Proteaceae, Leucadendron olens and Leucospermum hamatum , the succulent shrub, Lampranthus pauciflorus , and the herb, Linum villosum (Mucina & Rutherford 2006).

The conservation status of the type has been described as “vulnerable” (after the classification by Golding 2002) with large areas already transformed through afforestation and agricultural development (Mucina & Rutherford 2006, Lombard et al . 2005).

Tsitsikamma Sandstone Fynbos consists of mainly proteoid, restioid and ericoid fynbos with thicket in wetter areas, and is found on gentle to steep southern and northern slopes of the relatively low Tsitsikamma mountain range. Pierce (2003) further divides the type into Tsitsikamma Mountain Fynbos in the north and Tsitsikamma Plateau Fynbos to the south. Structurally, it can be described as medium dense, tall proteoid shrubland over a dense moderately tall, ericoid-leaved shrubland. As with South Outeniqua Sandstone Fynbos, it occurs on acidic lithosol soils derived from Ordovician sandstones of the Table Mountain Group, but plinthic catenas prominent (Mucina & Rutherford 2006). Fynbos islands are also found, but less widespread than for South Outeniqua Sandstone Fynbos. Floristic communities associated with the types, have been described by Southwood & De Lange (1984), Anon (1985), Hanekom et al . (1989), Pretorius et al . (1979). It has many species in common with South Outeniqua Sandstone Fynbos. Important species include the Proteaceae, Leucadendron conicum , L. eucalyptifolium , L. uliginosum subsp. glabratum and Protea mundii (tall shrubs), together with the Erica discolor variant speciosa and E. sparsa . Herbs include Commelina africana and Gazania krebsiana subsp. krebsiana , together with the Graminoids Restio triticeus and Tetraria capillacea . Endemic taxa include Aspalathus teres subsp. thodei , Erica trachysantha , E. zitzikammensis , Felicia tsitsikamae and Helichrysum outeniquense , with the conservation status of the type described as “vulnerable” (Mucina & Rutherford 2006).

North Outeniqua Sandstone Fynbos has a limited distribution in the park, largely restricted to gentle to steep north-facing slopes of the Outeniqua Mountains on the northern boundary of the park. It also occurs on acidic lithosol soils derived from Ordovician sandstones of the Table Mountain Group, with the Glenrosa and Mispah soil form prominent. It is dominated by restioid and proteoid Fynbos, with ericaceous and asteraceous fynbos widespread at higher and lower altitudes respectively. It has been described as tall, open to medium dense, medium tall shrub or restioid undertorey, with scattered, emergent, tall Proteaceae shrubs. Studies on floristic communities associated wit this type, has been conducted by Bond (1981). Important taxa include the small tree Protea nitida , the tall shrub Aspalathus sceptrum-aureum , the low shrubs Erica brachycentra , and the Graminoid, Heteropogon contortus . Species common with South Outeniqua Sandstone Fynbos and Tsitsikamma Sandstone Fynbos, include e.g. Leucadendron eucalyptifolium and Protea neriifolia . A number of endemic species have been recorded for the type, which include the shrubs Paranomus longicaulis and Aspalathus glabrescens , and the herbs, Oxalis attaquana and Haworthia outeniquensis . Its conservation status has been described as “least threatened” (Mucina & Rutherford 2006).

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Sand Fynbos

Two types of Sand Fynbos as described by Mucina & Rutherford (2006) have been recorded for the Park. Small pockets of Knysna Sand Fynbos are found along the Harkerville coast, while a small area of Southern Cape Dune Fynbos occurs in the Wilderness Section of the park.

Knysna Sand Fynbos are generally found on undulating hills and moderately undulating plains, and are described as dense, moderately tall, microphyllous shrubland. It is underlain by deep, acid Tertiary sand, forming regic sands and soils of the Lamotte form. Important species associated with this type, some also typical of Sandstone Fynbos, include the tall shrubs Cliffortia linearifolia , Leucadendron eucalyptifolium and Metalasia densa , the low shrubs Anthospermum aethiopicum and Berzelia intermedia , the herbs, Geranium incanum and Helichrysum felinum , and the Graminoids, Aristida junciformis subsp. galpinii and Brachiaria serrata (Mucina & Rutherford 2006). Vegetation units that form part of Knysna Sand Fynbos, have also been described by Anon (2005), who distinguish between Wet and Dry Sand Fynbos. Knysna Sand Fynbos has been rated by Mucina & Rutherford (2006) as “endangered” and very poorly researched. Lombard et al (2005) who consider it as almost totally transformed, close to extinction and urgently in need of protection.

Southern Cape Dune Fynbos is generally found on stabilised old calcareous or neutral dunes built with deep sand, often with steep slopes, but has a limited distribution in the park. It is dominated by sclerophyllous shrubs with a rich restio understorey. Dominant tall shrubs and small trees include Olea exasperate , Passerina corymbosa and number of Rhus species (e.g. R. crenata , R. glauca and R. lucida ). Smaller shrubs include Metalasia muricata , Phylica litoralis and Passerina rigida . Herbs include Satyrium princeps and Cyrtanthus loddigesianus , with Ischyrolepis eleocharis and Ehrharta calycina some of the dominant Graminoids (Mucina & Rutherford 2006). The type and different floristic communities associated with it are described in more detail by Van der Merwe (1976) and Anon (2005).

Although the conservation status of the type has been describes as “least threatened” it hosts endemic species such as Erica chloroloma , Lampranthus algoensis and Pentaschistis barbata subsp. orientalis . Aspalathus cliffortiifolia is also associated with the type, but is regarded as possibly extinct. Lombard et al . (2005), consider the type as heavily transformed.

Shale Fynbos

In the park, this type is represented by small patches of Garden Route Shale Fynbos found on the coastal foothills of the Outeniqua and Tsitsikamma Mountains. It consists of dense proteoid and ericaceous fynbos in wetter areas and more graminoid on drier sites. It is found on moist, clay- loam, prismacutanic and pedocutanic soils derived from the Caimans Group. With most of the shales covered by afromontane forest, Garden Route Shale Fynbos is found in areas regularly exposed to fires that disallow for forest development. Important taxa include the shrubs

management plan GRNP Appendices 15

Leucadendron eucalyptifolium , Protea aurea subsp. aurea and Cliffortia ruscifolia , the herbs Helichrysum felinum and Crassula orbicularis , and the graminoiods, Ischyrolepis sieberi and Aristida junciformis subsp. galpinii (Mucina & Rutherford 2006). Further species associated with Garden Route Shale Fynbos are provided by Anon (2004). It is a poorly studied vegetation type, rated as endangered, and hosts the endemic species Cyphia georgica , Disa newdigateae and Gladiolus roseovenosus .

Fynbos Shale Band Vegetation

In the park, this type is represented by Eastern Coastal Shale Band Vegetation . As the name indicates, it occurs on narrow, linear shale bands on clayey soils derived from the Cedarberg Formation. Important taxa include the widespread Leucadendron eucalyptifolium and Protea neriifolia , as well as L. salignum and Leucospermum cuneiforme . The type has been classified as “endangered” with the park regarded to make an important future contribution towards its protection (Mucina & Rutherford 2006).

Granite Fynbos

This type is represented by small areas of Garden Route Granite Fynbos in the lowland areas in the western part of the park (Wilderness area). It is dominated by proteoid and graminoid fynbos, on deep, prismacutanic- and pedocutanic-dominated soils, underlain by George Batholith of the Cape Granite suite. Important species include the shrubs Passerina corymbosa , Protea coronata and P. lanceolata , and the graminoids, Tetraria cuspidata and Brachiairia serrata . Garden Route Granite Fynbos has been classified as “endangered” with large areas already transformed for cultivation (Mucina & Rutherford 2006).

In addition to the “mainland” fynbos, there are a large number of smaller fynbos patches or “fynbos islands” of varying sizes. Most of these are natural in origin, attributed to the isolation of fynbos by forest expansion from a few centuries to several thousand years ago (Bond et al. 1988, Midgley & Bond 1990) and maintained by natural fire patterns (Geldenhuys 1994). Floristically they largely consist of South Outeniqua Sandstone Fynbos and Tsitsikamma Sandstone Fynbos, as well as smaller areas of lowland fynbos i.e. Knysna Sand Fynbos and Garden Route Shale Fynbos.

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APPENDIX D

FAUNA OF THE GRNP

ZOOPLANKTON

The highest diversity of species in Touw River Estuary has been recorded close to the mouth, with the highest daytime number of individuals in the Touw system recorded from Eilandvlei (14 641 indiv. m -3) and Rondevlei (13 108 indiv. m -3) (Coetzee 1983). The highest mean daytime standing crop has been recorded in Langvlei (17 mg m -3), followed by Rondevlei (15 mg m -3) and Eilandvlei (6 mg m -3) (Coetzee 1983). The lowest daytime planktonic standing crop in the Wilderness Lakes was recorded over deep areas in Swartvlei Lake, where bottom waters were deoxygenated (Grindley 1981; Grindley & Wooldridge 1973).

Zooplankton communities in the Swartvlei system consist primarily of estuarine species, with 45 forms having been recorded by Grindley & Wooldridge (1973). Coetzee (1981a, 1983) described two zooplankton communities during daytime, namely a mixolimnion community occurring mainly under aerobic conditions, and a monimolimnion community occurring primarily under anaerobic conditions and in the presence of H 2S. This division may be an artefact of sampling technique (Coetzee 1981a). Coetzee (1978 1981a 1983) found that at night there was a general upward migration in the water column of zooplankton and other benthic organisms, whereas Grindley & Wooldridge (1973) observed vertical diel migration only in Acartia spp, Halicyclops spp. and decapod larvae.

A comprehensive description of zooplankton communities in the Knysna estuary is given by Grindley (1985). Plankton in the channel north of Thesens Island is dominated by true estuarine species (Grindley 1985), with few freshwater species entering the estuary (Le Roi Le Riche & Hey 1947). A relatively low abundance of planktivores in Knysna Estuary suggests that plankton is not a major source of food in the Knysna estuary (Day 1967). The penetration of sea water into the estuary on each high-tide is thought to limit the survival of true estuarine species to areas where the residence time of water is sufficiently long to enable them to complete their life cycles (Grindley 1985). During cold upwelling events various deep ocean species which do not usually form part of the estuarine or neritic marine plankton community may enter the estuary. Morant and Bickerton (1983) listed more than 25 taxa of zooplankton recorded from the Groot River (West) estuary

management plan GRNP Appendices 17

INVERTEBRATES

Estuarine aquatic invertebrates

Benthic invertebrates in the open areas of Swartvlei Lake have been sampled by Boltt (1973) who collected 14 predominantly estuarine species. The dominant species include the amphipoda Corophium triaenonys, Cyathura estuaria, Grandidierella lignorum and Melita zeylenica , isopoda Exosphaeroma hylecoetes and Cyathura estuaria , tanaid Aspeudes digitalis , bivalve Musculus virgiliae , polychaete Ficopomatus enigmatica , crab Hymenosoma orbiculare and prawn Callianassa kraussi (Davies 1982). Most of the zoobenthos occur on submerged aquatic plants (Davies 1982), with the mean biomass on Potamogeton pectinatus in the Touw system calculated during 1979 and 1980 to be 77.6 g dry mass m -2, compared to the 12.7 g dry mass m -2 recorded in sediments (Davies 1982).

Abundant species of benthic invertebrates in Swartvlei Estuary include Melita zeylinaca , Loripes clauses , Natica tecta and Palaemon pacificus in vegetated areas (principally Zostera capensis ), whereas those of bare sand areas are Urothoe pulchella , Callianassa kraussii , Iphinoe truncate and Pontogeloides latipes (Whitfield 1989e). The supratidal invertebrate community is dominated by Orchestia spp. which feed on wrack detritus. Diet analysis of intertidal and infratidal zoobenthos in Swartvlei Estuary revealed that most invertebrate species feed predominantly on detritus and associated micro-organisms. Filamentous algae and diatoms dominate the gut content of only three out of 18 macrobenthic species, and living Zostera is not an important food item for any invertebrate species (Whitfield 1989e).

The benthic macrofauna in Knysna Estuary includes approximately 310 species (Day 1981). The ecology of some species, such as the endemic mudsnail Hydrobia knysnaensis (cf. Barnes 2004) have been the focus of detailed studies. Day et al . (1952) demonstrated that the diversity of aquatic invertebrates progressively declines upstream of The Heads. Furthermore the percentage of typical seashore species decreases progressively up the estuary, with none occurring at Charlesford Rapids (Day et al . 1952). Estuarine species are widespread in the estuary, whereas “brack-water species” favour oligohaline conditions at Charlesford Rapids (Day et al. 1952). Studies similar to those reported by Day et al (1952) were also undertaken in 1997 (Allanson et al 2000b). Comparisons of data from different surveys showed that there was no difference between species richness of transects reported in 1952 and 1997. A significant increases was however recorded in species diversity (Shannon-Weiner Index) in quantitatively sampled sediments in the Zostera zone which was though to be due to intermittent increases in fluvial derived suspensoids in the water column (Allanson et al. 2000b). It has been suggested that an increase in silts and clays (<60 m) after settling has altered the quality of intertidal sediments, making them more suitable for increases in both the number of individuals of resident taxa as well as new taxa (Allanson et al . 2000b).

management plan GRNP Appendices 18

The mudprawn Upogebia africana is abundant and is the most widely used bait species in Knysna Estuary (Patterson 1986). Upogebia africana occupy 62% of the available intertidal zone, with density (74-76 m -2) and biomass (26-27 g m -2 dry weight) usually greatest in the Spartina and lower Zostera zones (Hodgson et al. 2000). In 1995 the Invertebrate Reserve was found to have a low density and biomass of U. africana , (11.7 m -2; 3.9 g m -2) whereas a relatively inaccessible mudbank (Oyster Bank) in the middle reaches of the estuary had a much larger population (176 m - 2; 6.5 g m -2) (Hodgson et al . 2000 ). This difference could be a result of more favourable habitat at the Oyster Bank (e.g. less variable temperature and salinity) and/or the inaccessibility of the site to bait collectors (Hodgson et al . 2000).

The critically endangered false limpet Siphonaria compressa , originally described by Allanson in 1958 from the Langebaan Lagoon, has been found living in intertidal eelgrass meadows at Bollard Bay, Leisure Isle (Allanson & Herbert 2005). This represents only the second known site of occurrence of the species. In Langebaan Lagoon it is confined to the lower edge of the eelgrass beds (Angel et al. 2006). Rarity of S. compressa and its endangered status seem dictated by its extremely narrow and temporally changeable habitat range (Angel et al. 2006). The Knysna population is considered to be viable (Allanson & Herbert 2005).

Three species of indigenous oyster species occur in Knysna Estuary, namely South African Oyster (Crassostrea margaritacea ), 'weed-oyster' ( Ostrea algoensis ) and Red Oyster ( Ostrea atherstonei ) (Grindley 1985). The Pacific Oyster ( Crassostrea gigas ) was introduced in 1973 and today forms the basis of a flourishing local industry.

Freshwater aquatic invertebrates

Early studies of aquatic invertebrates in southern Cape rivers were undertaken by Harrison & Agnew (1962). Filmalter & O’Keeffe (1997) provide a comprehensive list of invertebrates in rivers flowing into the Wilderness lakes. Biotic index values occasionally exceeded 2 in the Diep River indicating slightly enriched waters. In the Hoëkraal, Karatara, Duiwe and Touw river biotic index values were always below 2.

Aquatic invertebrate surveys in the Salt River showed that in the four orders of insects examined in detail (Plecoptera, Ephemeroptera, Trichoptera and Diptera) there were 13 undescribed species, as well as three possible new genera, and some remarkable range extension of certain species. The river also produced the richest known diversity of species in the mayfly family Teloganodidae for Africa (De Moor et al. 2004). Water abstraction from the catchment of the Salt River is the most serious threat to the continued existence of these macroinvertebrate communities, and reduced flow volumes can exacerbate or result in a multitude of detrimental impacts (De Moor et al. 2004).

Terrestrial Invertebrates

management plan GRNP Appendices 19

Insects are the most diverse group of organisms on earth (Picker et al. 2003). The Garden Route National Park’s position within two vegetation biomes, one a floral kingdom and biodiversity hotspot, contributes to a range of habitats in diverse successional stages that no doubt harbor rich invertebrate species diversity. Even though terrestrial invertebrates on earth exponentially outnumber the vertebrates, they are relatively poorly understood on a global level, a situation that applies to the GRNP and surrounding area. Van der Merwe (2002) mentions that a multitude of insects, spiders, millipedes, centipedes, worms and snails populate every niche between the forest floor and the canopy and yet the forests still harbor thousands of invertebrate species unknown to man. Cameron (1982) states that although it may be true that forests support a small animal fauna if only the vertebrates are being considered, it is also true that the greatest variety and number of invertebrates are found in the forest habitat. Many animal species are specifically adapted to the characteristic ecological processes of the forest environment and of different forest development stages (Geldenhuys, 1998). Several insect genera are confined to the forest biome and are host specific to certain plant species or genera (Geertsema 1964; Scholtz & Holme, 1985; Swain & Prinsloo, 1986).

Most species of the Myriapoda , Arachnida , Anychophora , terrestrial Crustacea and wingless insects ( Apterygotes ) are found in forests. Cameron (1982) mentioned that this cryptofauna is remarkably homogenous in South African forests, especially in orders and families represented. The terrestrial Amphipoda and Isopoda , the terrestrial planarians, the Onychophora , the Palpigradi , Opiliones and Acarina among the Arachnida , the Pauropoda , Symphyla and Polydesmoidea among the Myriapods and practically all the Apterygote insects are shade and moisture loving orders typical of the cryptic life of the humus layer of the forest floor.

While a few specialist invertebrate surveys have identified several species (Appendix C), insect collections to date have been uncoordinated and are not representative of the entire GRNP as large parts of the region have never been surveyed for invertebrates, and existing data consists largely of scattered records and species descriptions for individual taxa. The diversity, biology, ecology and conservation needs and status of the fauna are poorly known. Conservation of the Park’s invertebrates is critical because of their roles in ecosystem functioning, the dependence of many vertebrate species on invertebrates as a source of food, and also because of the potentially large numbers of species and genera endemic to the region. Hamer (2005) mentioned that endemics are more likely to be represented by less mobile, moisture dependent, forest dwelling taxa such as millipedes, earthworms, spiders and mollusks. Nevertheless, it is assumed that the invertebrate species identified in these surveys are representative of the invertebrates inhabiting similar habitat within the GRNP.

Crewe (1984) surveyed insects in the Harkerville forest and fynbos areas. These surveys revealed that numerous insects belonging to most of the insect families occur within the forests and the area is described as being particularly rich in cryptic insect fauna.

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Grindley (1985) collected terrestrial invertebrates in the Knysna Lake Area. Insects collected included chironomid midges, dragonflies, mayflies, kelp flies, staphylinid and other beetles, house flies and water boatmen.

Breytenbach (1988) investigated ant-plant mutualistic interactions in the Outeniqua Mountains near George and Koen & Breytenbach (1988) researched the species richness of ant fauna in fynbos and forest habitats in the Gouna and Diepwalle areas. No significant difference in ant species richness was found between the two undisturbed habitat types, and the only two species common to both are Acantholepis capensis and Camponotus maculatus . Ant species richness in the Afromontane forest decreased from the drier to the wetter sites and that the degree of Hakea sericea infestation in the fynbos negatively affected ant species richness. They also indicated that the aggressive alien argentine ant, Iridomyrmex humilis , has not invaded natural areas to any great extent. Ant species were also collected on fynbos islands, mainland and forest in the Diepwalle area (Midgley and Bond, 1990).

Marais (1991) surveyed and compiled a checklist of the butterflies of the Tsitsikamma National Park, which was supplemented by Butler and Terblanche (1997). 41 butterfly species/subspecies have been recorded in the De Vasselot section of the park. Edge (pers comm.) indicated that the forest butterfly species are well surveyed and that about 13 species of butterfly are listed. Three species ( Charaxes karkloof trimenii , Symothoe althemeda and Charaxes xiphares ) are endemic to the region. Apart from being endemic, Charaxes karkloof trimenii was only discovered within the last decade. Edge also mentioned that the mountain catchment areas are poorly surveyed and that the possibility of discovering new species is good.

A moth survey was conducted in the Diepwalle indigenous forest and fynbos area by Kroon (1998). 232 moth species were identified, 9 butterfly and one beetle species. Minuscule moths of the genus’s Epiplema , Heteroplema and Monoplema were also identified.

(Geldenhuys 1993c) found that occasional insect outbreaks occur, especially when there is a drought season following a wet period. These outbreaks are probably a normal cyclic event and as such are unlikely to pose a major threat to forests.

An invertebrate of significance found in the forest areas is Peripatus . It is an ancient life form that is believed to have survived unaltered for 500 million years and can be considered as living fossils dating back to the Cambrian period (Hey, 1973 cited in Cameron, 1982). Its kidney and blood system resembles those of worms and the breathing system those of insects. It is also found in the forests of South America, Australia and Asia, providing support for the theory of continental drift. Peripatus is protected under the Provincial Nature Conservation Ordinance (CEN Integrated Environmental Management Unit, 1999; Durrheim, 2002). Another discovery supporting the continental drift theory is the close resemblance between the swift moth ( Phalaena venus ), endemic to the southern Cape forests and the Australian swifts. This moth occurs in close association with its larval host tree, the pink blossom tree ( Virgilia divaricata ). Almost every sizeable pink blossom tree is infested by the

management plan GRNP Appendices 21 larvae, which bore into the roots and the base of the trees. Some trees die as a result. However, the moth itself is seriously threatened by the spread of the exotic Argentine ant.

Members of Onycophora , a rare, primitive Arthropod phylum, are relatively abundant in the Harkerville area and were collected by researchers from the United Kingdom (Vermeulen et al. , 1995). A number of apparently new species of Drosophilae were also collected in the area (Vermeulen et al ., 1995).

A rare butterfly species, Chrysoritis cottrelli , has been found at Spitskop, but was last seen in 1975 (Muller, 1990). Its larvae live on bietou ( Chrysanthemoides monilifera ), their host plant. Since this plant species occurs in the Ysternek Nature Reserve area, it is possible that this small orange butterfly may still be found there (Durrheim, 1993).

The Brenton Blue butterfly ( Orachrysops niobe ), arguably the best known terrestrial invertebrate found in the Knysna Lake area became the focus of conservation efforts following the threatened destruction of the habitat of the last known population at Brenton-on-Sea by a housing development (Steenkamp & Steyn, 1999). Assessments were undertaken of the life-history and ecology of this species, as described by Edge & Pringle (1996) and Williams (1996) to enable effective conservation of remaining populations. The asteraceous fynbos where O. niobe occurs is characterised by a great diversity of shrubs, herbs and graminoids, with a successional gradient to thicket where Pterocelastrus tricuspidatus is dominant (Lubke et al. 2003). The eggs of the butterfly are laid on the lower side of the leaves of Indigofera erecta. The butterfly appears to require a very specific stage in the coastal fynbos vegetation for breeding sites, and fire probably maintains this mosaic of vegetation types. The vegetation should best be maintained with fire, burning at intervals of more than 10 years (Lubke et al. 2003).

Six mosquito species of the genera Anopheles , Aedes , and Culex have been collected in the Wilderness area (Russell & Russell 1997). Culex salisburiensis has been collected at only two localities, one of which is Wilderness.

Endangered species that possibly occur (Picker et al , 2004) in the GRNP include all the Colophon (Stag Beetles) spp. The genus is highly sought after by collectors and is thus listed as a red data book genus.

Protected invertebrate species that possibly occur (Picker et al , 2004) in the GRNP include Hadogenes spp, Harpactira spp, Opithacanthus sp p – all species, Opistophthalmus spp – all species, Pterinochilus spp – all species and Manticora spp – all species. These are indigenous species of high conservation value or national importance that require national protection.

A trade in butterflies and insects from forests is conducted on a legal and illegal basis. A large illicit trade exists for Cicadas. Two species of cicada are known to occur in the forests of the GRNP.

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Invertebrates play critical roles in the functioning of all ecosystems as they are responsible for maintaining soil fertility, for waste disposal, water purification, pest control and pollination. Few studies quantifying the contribution of invertebrates to these processes have been performed in South Africa, but internationally the complexity of the invertebrate interactions required to sustain ecosystems and even in influencing the structure of plant communities is becoming increasingly evident (Hamer 2005). Numerous invertebrates, for example earthworms, are considered to be keystone species by recycling large amounts of plant biomass into the soil and keep the soil porous with their tunneling, allowing water to infiltrate the soil profile. Pollination of a vast proportion of flowering plants, including endemics, is dependant on a range of insect groups, such as bees, wasps, flies, and beetles. In some cases the survival of locally endemic plant species is linked to pollination by a single insect species.

FISHES

Estuarine fishes

Accounts of fishes in the Swartvlei system are given by Kok & Whitfield (1986) and Russell (1996a), and in the Touw system by Hall (1985a, 1985b), Hall et al . (1987) and Russell (1996a), who demonstrate that communities are typically dominated by juvenile marine species. Both Hall (1985a) and Russell (1996a) found that the number of species in the Touw system declines the further removed a waterbody is from the sea. Hall et al . (1987) have suggested that several fish species may experience difficulty in negotiating the inter-connecting channels between lakes. Thus the closed phase of the Touw system, when water levels are higher, may be important to fish migration as it would reduce the restrictions on distribution imposed by interconnecting channels.

Patterns of recruitment of juvenile fishes into Swartvlei Estuary have been studied by Kok (1981) and Whitfield (1989a, 1989d) who found strong summer recruitment for most species, with optimum recruitment periods for most dominant species being from October to February. This is similar to the optimal period of September to March given by Kok et al . (1981). An extended spawning period is thought to be an adaptive strategy whereby the period of potential juvenile recruitment is prolonged, in effect creating a buffer against failure of recruitment as a result of adverse marine or estuarine conditions (Whitfield & Kok, 1992). The importance of the surf zone for estuarine associated fishes has been demonstrated by Whitfield (1989c, 1989d), with larvae and postlarvae of 16 families identified from the surf zone off Swartvlei Estuary. Densities peak during summer and when the estuary mouth is open, with concentrations highest closest to the estuary mouth. Time of day and tidal phase may be important in governing ichthyoplankton abundances in the surf zone (Whitfield 1989c). The effect of prolonged closed phases on the recruitment of fishes in the Wilderness lakes was evaluated by Russell (1996a). The importance of also having a closed phase in the Wilderness Lakes estuaries to provide an ideal nursery habitat, in terms of nutrition for juvenile marine fishes is emphasised by Kok & Whitfield (1986). Whitfield (1992) has

management plan GRNP Appendices 23 demonstrated that recruitment of juvenile fishes form the surf zone into a closed estuary can occur during high seas when the sand bar at the estuary mouth is overtopped.

Extensive studies have been undertaken of the feeding habits of the dominant fishes in the Wilderness lakes, including the leervis Lichia amia (Smale & Kok 1983; Coetzee 1982a; Whitfield 1988a), Knysna halfbeak Hyporhamphus knysnaensis (Coetzee 1981b; Whitfield 1988a), sea barbel Galeichthys feliceps (Coetzee & Pool 1984), Cape sivlerside Atherina breviceps and estuarine roundherring Gilchristella aestuaria (Coetzee 1982b; Whitfield 1988a), Cape stumpnose Rhabdosargus holubi and Cape moony Monodactylus falciformis (Whitfield 1988a), as well as the strepie Salpa salpa , blacktail Diplodus sargus , southern mullet Liza richardsonii , groovy mullet Liza dumerilii , flathead mullet Mugil cephalus , white steenbras Lithoglanis lithoglanis , Sea catfish Galeichthys feliceps , Kob Argyrosomus hololepidotus , super klipfish Clinus superciliosus , prison goby Caffrogobius multifasciatus , barehead goby Caffrogobius nudiceps , Knysna sandgoby Psammogobius knysnaensis and longsnout pipefish Syngnathus acus (Whitfield 1988a). Based on these studies, estuarine fish can be grouped into five feeding categories, namely detrivores (e.g. L. richardsonii, L. dumerilii, M. cephalus ), herbivores (e.g. S. salpa ), omnivores (e.g. R. holubi, D. sargus ), carnivores (e.g. P. commersonnii, M. falciformis ) and piscivores (e.g. L. amia, A. hololepidotus ). Whitfield (1993) estimated the biomass of the littoral fish community in Swartvlei Lake during 1980 to be 12.4 g m -2 wet weight, with detrivorous species contributing 3.2 g m -2 , zoobenthic consumers 2.8 g m -2 , piscivorous species 2.3 g m -2, herbivorous/epifaunal consumers 2.7 g m -2 and zooplanktivorous consumers 1.4 g m -2. It was concluded that estuarine fish biomasses do not exceed those of productive freshwater or marine environments. Changes in fish communities as a result of alteration in the food resource was observed in the early 1980s where the senescence of submerged macrophyte beds over a three year period resulted in a 60% decline in primary production, a 74% decline in littoral invertebrate biomass, and 54% decline in the abundance of the fishes M. falciformis and R. holubi associated with the macrophytes (Whitfield 1984). Furthermore, an increase in epipsammic micro-algae production in the littoral during the macrophyte senescence phase resulted in an increase in the abundance of the family Mugilidae (Whitfield 1986). However, the resilience of estuarine fish communities to major habitat and food resource changes was also demonstrated by all fish species recorded in Swartvlei Lake at the beginning of the macrophyte senescence phase still being present after three years of extensive habitat change (Whitfield 1986).

Detailed studies of the age structure, growth and reproductive biology of Atherina breviceps and Gilchristella aestuaria in Swartvlei Lake have been made by Ratte (1989). It was observed that 0- year-old A. breviceps and 2-year-old G. aestuaria dominated catches in Swartvlei Lake. Annual production of A. breviceps was 10 tons, and G. aestuaria was 5 tons. Ages attained were three years for A. breviceps , and seven years for G. aestuaria . The main spawning period for A. breviceps is spring-autumn, whereas G. aestuaria has continuous spawning.

The Knysna seahorse (Hippocampus capensis ), classified as Endangered (Hilton-Taylor 2000), is widespread in Swartvlei and Knysna estuaries though not abundant. Lockyear et al. (2006) estimated the adult population density in Swartvlei Estuary to be 0.26 individuals m -2 (Jan-Feb

management plan GRNP Appendices 24

2002) and 0.04 individuals m -2 (Dec 2002- Jan 2003). Bell et al. (2003) estimated the adult population in Knysna Estuary to be 89 000 individuals (range 30 000 to 148 000: 95% confidence interval), and Lockyear et al. (2006) estimated the adult population to be 62 000 individuals in the subtidal areas which were estimated to be 5.4 x10 6m2 or 33% of the surface area of the estuary. Bell et al. (2003) found H. capensis in Knysna Estuary most frequently in low density vegetation stands ( ≤ 20% cover) and grasping Z. capensis . Seahorse density is not otherwise correlated with habitat type or depth. Toefy (2000) using conventional meristics, morphometric work and limited sequencing of the mitochondrial cytochrome b gene of H. capensis , found that a certain amount of variation existed between the Knysna and Swartvlei populations, and recommended that mixing of the two populations should be avoided. An alternative management proposal was forwarded by Teske et al . (2003) who investigated the evolutionary history of H. capensis using 138 mitocondrial DNA control region sequences. Based on haplotype frequency distribution it was concluded that the three known assemblages (Knysna, Swartvlei, Keurbooms) constitute distinct management units, though it couldn’t be concluded that they are evolving relatively independently under different stochastic processes. The absence of distinctive monophyletic clades of haplotypes unique to individual populations suggested that at this stage there is little reason to discourage the translocation of seahorses amongst the different estuaries. The age of the Knysna population has been estimated to between 46 000 and 486 000 years (late Pleistocene) (Teske et al . 2003), which suggests that apart from being geographically isolated from its sister species and living in a habitat that is likely to be inhospitable to other seahorses because of unstable physical and chemical conditions, H. capensis may also be phylogenetically distinct (Teske et al . 2003). Consequently, the high conservation status of this species seems justified. Breaching of Swartvlei Estuary has been observed to result in the mortality of Knysna seahorses ( Hippocampus capensis ) with in 1985 in excess of 100 individuals washed up on the banks of the estuary, presumably killed by low salinity (<15 g kg -1) (A. Whitfield pers comm.) which has been shown to be physiologically stressful for the species (Riley 1986). Most subsequent die-off’s have involved less than100 individuals (SANParks unpublished data) though the largest die-off occurred on the 18 February 1991, with mortality estimated to have exceeded 3000 individuals, resulting from high water temperatures (32°C) which occurred in the shallow marginal areas of the Swartvlei Estuary following an extended period of hot weather (Russell 1994).

In excess of 200 species of fish have been recorded in the Knysna Estuary (Bulpin 1978). A complete species list is given in Grindley (1976), with the more common species listed in Grindley (1985). The permanently open estuary enables free access to typical marine species, with the result that there are many records of species which do not normally occur in estuaries (Grindley 1985). What is thought to be a new species of goby from the Knysna river Gobius maxillaris sp. n. was described by Davies (1948). No specimens of this species have been recorded in the Knysna estuary, or any other estuary, since this time. A rare estuarine goby, Pandaka silvana , is endemic to Knysna Estuary (Penrith & Penrith 1972). Le Quesne (2000) investigated the usage by fish of intertidal marshes dominated by Juncus kraussii in the upper reaches of Knysna Estuary. Twenty five, predominantly euryhaline marine species were recorded, which were dominated numerically by juvenile (<30mm) Mugilidae, Atherina breviceps and Liza richardonii , and in terms of biomass by L. richardsonii , Lithognathus lithognathus , Pomadasys commersonnii and Rhabdosargus

management plan GRNP Appendices 25 holubi . Intertidal marshes are argued to provide areas of refuge from predators and feeding opportunities, with the nine most dominant taxa actively feeding in marsh areas. As such transient marsh nekton may be an important conduit for marsh production into the estuarine and coastal ecosystem.

Harrison et al. (1996) recorded 15 fish species in the major estuaries of the Tsitsikamma region. Three of the species were estuarine-dependent, the others were marine species, having various degrees of dependence upon estuaries during the juvenile phase of their life cycle.

Marine fishes

A rich diversity of fish is found in the marine section of the park, with a total 202 fish species from 84 families recorded (Buxton and Smale 1984; Burger 1991; Wood et al. 2000) The size and species composition of fish differ between shallow and deep reefs in the park (Buxton and Smale 1984), and species diversity and richness tends to increase with depth (Burger 1990).

The biology and life history changes of 10 commercial or recreationally exploited marine fish species conserved in the park have been studied. These include dageraad, roman (Buxton, 1987 1989, 1990, 1992, 1993), red steenbras (Smale 1988; Smale & Punt 1991), blue hottentot (Buxton & Clarke 1986), hottentot (Buxton & Clarke 1992), musselcracker (Buxton & Clarke 1991), poenskop (Buxton & Clarke 1989), blacktail, zebra ( Mann & Buxton 1992, 1998) and carpenter (Brouwer & Griffiths 2004; 2005; Brouwer et al. 2003).

Most (c 80 %) of the above fish species are slow growing, long-lived species (>20 years), and many have a high degree of residency (Buxton 1987; Cowley 2000; Cowley et al. 2002; Brouwer et al . 2003), which makes them vulnerable to over-exploitation. A point highlighted by the fact that the catch rates during research studies in the park were considerably higher than those recorded for fishers in nearby exploited areas (Buxton & Smale 1989; Brouwer & Buxton 2002; Brouwer 1999; Cowley et al . 2002; Cowley 2000).

Freshwater fishes

Nine fish species have been recorded in the Duiwe and Touw rivers (Russell 1999b) three of which are alien. A further one indigenous species, two translocated species and five estuarine species could potentially occur in these rivers (Russell 1999b). Freshwater fishes recorded in the Knysna River include “ Barbus monodactylus ” (probably Pseudobarbus afer ) “ Sandelia spp.” (probably Sandelia capensis ) and “springer” (probably Myxus capensis ) (Le Roi Le Riche & Hey 1947). Both black bass and trout species have been introduced in the past though appear not to have become established (Le Roi Le Riche & Hey 1947). Le Quesne (2000) reported collecting the alien Gambusia affinis in intertidal marshes in the upper reaches of the estuary. The fresh water streams in the Tsitsikamma region are generally impoverished (Smith & Smith 1966). Russell (2002) sampled six river systems in this region and recorded four indigenous freshwater species

management plan GRNP Appendices 26

(Pseudobarbus afer , Pseudobarbus tenuis , Sandelia capensis and Anguilla mossambica ) and one alien ( Micropterus salmoides ).

REPTILES

Introduction

The Garden Route National Park’s (GRNP) location within the fynbos (Floral Kingdom and Biodiversity hotspot) and rare forest biome, the inclusion of a marine and unique wetland/estuarine lake system and the rugged mountain and coastal landscape provides diverse habitat in varied successional stages that consequently contribute to a reasonably diverse reptile fauna. Moreover, the GRNP forms the eastern limit of a distinct zoogeographic zone, called the Cape Faunal Centre where the fauna gradually changes to include more Ethiopian elements characteristic of much of Africa. Unfortunately, little is known of the reptiles inhabiting the National Parks of the eastern and southern Cape (Branch, 1981, 1983).

Species Richness/Diversity

Systematic knowledge of the Western Cape Province’s (WCP) reptiles is still in an alpha taxonomic stage with new species still being discovered and described (Turner et. al 2007). In the last five years in the WCP, at least nine new species have been discovered and one has been described (Bauer et al. 2003). This mirrors the situation in southern Africa generally where current estimates put the total number of reptile species at 520 or more (Branch 2006), which represents a dramatic increase from the last published total of 397 (Branch 1998).

The Garden Route National Park is considered to fall within one of the eight major centres of herpetofauna diversity in southern Africa (Branch, 1998) and forms part of the mountainous region stretching from the Cape in the south northwards to tropical Africa that is considered to function as a corridor along which various faunal taxa can move.

Reptile checklists were derived from Von Breitenbach (1974), unpublished records from Cape Nature Conservation and Grindley (1985). Checklists were compiled by Branch and Hanekom (1987); Whitfield ( et al . 1983) and unpublished (SANParks) records of occurrence of species. All the reptiles that might occur inside the reserve (Von Breitenbach, 1974; Branch, 1988) are listed in Appendix C.

In total, the world contains more than 6 550 species of reptiles that are arranged in 900 genera in 48 families. Branch (1998) states that southern Africa perhaps has the highest reptile diversity on mainland Africa, and that the lizard fauna is by far the richest and most diverse. This is particularly the case among the geckos, skinks and girdled lizards. South Africa has a total number of 411+ reptile species (Turner et al ., 2007), 6.27% of the world total and 148 (36% of SA) species have

management plan GRNP Appendices 27 been recorded in the WCP. Fifty-six species of reptile comprising two orders and seventeen families most probably occur within the GRNP, which means that only 13.6% of the species occurring in South Africa also occur in the Park. Similarly only 38% of reptiles that occur the WCP occur in the GRNP and this total comprises 23 (41% of the GRNP total) lizard, 25 (45%) snake, three terrestrial tortoises (5.4%), four marine turtles (7.1%) and one (1.8%) freshwater terrapin species. Forty-six species (82%) have been formally recorded within the Park boundaries (Appendix A). Five species (9%) are restricted to the marine ecosystem and are classified as vagrants. The larger of the two orders is Squamata that represents eight-six percent of the parks reptiles and is comprised of two sub-orders, thirteen families and forty-eight species. The second order, Chelonii, has one sub-order, four families and eight species.

The true species total is likely to be higher as the distribution ranges of number of species that could possibly occur within the Park end just short of Knysna. It is uncertain whether the leopard tortoise ( Geochelone pardalis ) occurs naturally or has been artificially introduced into the area.

Table 1. Orders, families and reptile species of the GRNP:

ORDER SUB-ORDER FAMILIES SPECIES REDLIST IUCN CITES Chelonii 1 4 8 4 4 7 Squamata 2 13 48 Null Null 5 3 17 56 4 4 12

Endemism

Thirty-six species have varying degrees of endemism (Table 2). Five species are Cape Floristic Kingdom endemics, namely Tetradactylus africanus, Tropidosaura gularis, Tropidosaura montana montana, Bradypodion damaranum and Cordylus coeruleopunctatus. Three species are Cape Floristic Kingdom near-endemics, namely, Pseudocordylus microlepidotus microlepidotus , Afrogecko porphyreus and Homopus areolatus . Thirteen species are endemic to South Africa and fifteen species are southern Africa endemics. Apart from being CFR endemics, two species, Bradypodion damaranum (Knysna dwarf chameleon) and Cordylus coeruleopunctatus (blue- spotted girdled lizard) are entirely restricted to the southern Cape area (Branch and Hanekom 1987; Branch 1988). Another species, Hemachatus haemachatus (Rinkhals) although a southern Africa endemic, has relict population along south cape coast.

management plan GRNP Appendices 28

Table 2. Reptilian species endemic/near-endemic to the CF R, RSA and southern Africa.

Species C.F.R. C.F.R. RSA Southern endemic near- Africa endemic Tetradactylus africanus X Tropidosaura gularis X Tropidosaura montana montana X Bradypodion damaranum X Cordylus coeruleopunctatus X Pseudocordylus microlepidotus X microlepidotus X Afrogecko porphyreus X Homopus areolatus X Chamaesaura anguina anguina X Pachydactylus geitje X Tetradactylus seps X Tetradactylus tetradactylus X Nucras lalandii X Acontias meleagris meleagris X Mabuya homalocephala smithii X Duberria lutrix lutrix X Lamprophis aurora X Lamprophis inornatus X Philothamnus natalensis occidentalis X Homoroselaps lacteus X Cordylus cordylus Chersina angulata X Agama atra atra X Pachydactylus bibronii X Pachydactylus maculatus X Pedioplanis lineoocellata pulchella X Eremias lineocelata X Amplorhinus multimaculatus X Lycodonomorphus rufulus X Prosymna sundevallii sundevallii X Psammophis crucifer X Psammophylax rhombeatus X Hemachatus haemachatus X Naja nivea X Rhinotyphlops lalandei X Bitis atropos X The Tsitsikamma Mountains area (east) has also been identified as containing isolated populations of the endangered Smith's dwarf chameleon (Bradypodion taeniabronchum) and proper management of mountain fynbos habitats is important in ensuring the continued survival of suitable habitats for this taxon. (Baard et al . 1999).

management plan GRNP Appendices 29

In addition, sixteen species (35.7%) are range restricted. Eight species do not occur further east than Port Elizabeth and eight species have western distribution limits, two of which do not occur west of Riversdale, namely Causus rhombeatus and Philothamnus natalensis occidentalis. All four turtle species are vagrants to the park (Branch and Hanekom 1987; Branch 1988).

Conservation Status

The Park protects eleven species with varying degrees of conservation status (Table 3). The four marine turtle species, namely Caretta caretta, Chelonia mydas, Eretmochelys imbricata and Dermochelys coriacea that originate from one order and two families are listed as vulnerable in the South African red data book (Branch, 1988). Eretmochelys imbricata and Dermochelys coriacea are listed as critically endangered according to IUCN ratings. Both these species typically frequent warmer waters to the north and east of the WCP and do not breed on WCP shores. However, it is unknown to what extent they are making use of WCP waters as migratory routes or feeding sites. It is also unknown to what extent fishing activities in WCP marine waters affect these species. Several specimens, and mostly sub-adults, have washed up on WCP shores, usually following periods of rough seas or of significant upwelling events following strong off-shore winds causing ocean temperatures to drop significantly. Caretta caretta and Chelonia mydas are listed as endangered (Table 3). Seven species are listed as protected according to the NEMBA category (Table 3).

The Southern African Reptile Conservation Assessment (SARCA) was launched in May 2005 to begin a revision of the distribution and conservation status of Southern Africa’s reptile species, and will revise the outdated SA Red Data List for Reptiles and Amphibians (Branch 1988). It is extremely likely that more reptile species occurring within the GRNP will be on this new list as much more information and research are available on a number of species, including a number of recently described species that warrant conservation measures. For instance, a revision of the current status of CFR herpetofauna yielded a number of new, proposed IUCN Red List taxa (Baard et. al . 1999; Mace and Lande, 1994).

Table 3. Conservation status of reptiles that occur in the GRNP

Order Family Scientific English SARDB IUCN CIT NEMBA Name Name Status Status ES Category Chelonii Cheloniidae Caretta caretta Loggerhead Vulnerable Endangered App 1 Endangered turtle (A1abd) Chelonii Cheloniidae Chelonia mydas Green turtle Vulnerable Endangered App 1 Endangered (A2bd) Chelonii Cheloniidae Eretmochelys Hawksbill sea Vulnerable Critically App 1 Critically imbricata turtle Endangered Endangered (A1bd) Chelonii Dermochelyidae Dermochelys Leatherback Vulnerable Critically App 1 Critically

management plan GRNP Appendices 30

coriacea sea turtle Endangered Endangered (A1abd) Chelonii Testudinidae Chersina Angulate Null Null App 2 Protected angulata tortoise Chelonii Testudinidae Geochelone Leopard Null Null App 2 Protected pardalis tortoise Chelonii Testudinidae Homopus Parrot-beaked Null Null App 2 Protected areolatus tortoise Squamata Chamaeleonidae Bradypodion Knysna dwarf Null Null App 2 damaranum chameleon Squamata Cordylidae Cordylus Blue-spotted Null Null App 2 Protected coeruleopunctatu girdled lizard s Squamata Cordylidae Cordylus Cape girdled Null Null App 2 Protected cordylus lizard Squamata Cordylidae Pseudocordylus Cape crag Null Null App 2 Protected microlepidotus lizard microlepidotus

Twelve species (21%) are CITES listed (Table 8). The four marine turtles mentioned above are listed as Appendix I species and the eight remaining species fall in the Appendix II category. Thirty-three species (58.9%) are also listed in the WCP ordinance schedule (Appendix 1).

Ecological Studies

Ecological Knowledge of the reptiles of the GRNP is limited, consisting primarily of published and unpublished (SANParks) records of occurrence of species. The only paper produced on reptiles in the GRNP till date was by Branch and Hanekom (1987) in which an annotated checklist was produced comprising 38 species. The paper also discusses their ecology and zoogeographic affinities. They found that both the forest and coastal sections of the Park had very similar herpetofaunas but that the diversity of reptiles along the Tsitsikamma coast is greater than that of the forest section of the park. This is in accordance with Branch (1981) who indicated that few lizards, with the exception of chameleons, have specifically adapted to forest niches, which are generally too wet and cool and unsuitable for most reptiles.

Since the last reptile assessment was carried out 16 years ago and is very outdated, the South African Reptile Conservation Assessment (SARCA) was launched in 2005 with the intention of replacing the outdated South African redlist. This is a participatory project that relies heavily on people to record and report sightings. This project will be gathering data on reptiles up until the end of 2008 and it is important that Park staff participate actively to facilitate the conservation status of the Parks reptiles.

management plan GRNP Appendices 31

AMPHIBIANS

Introduction

The Garden Route National Park’s (GRNP) location within the fynbos (Floral Kingdom and Biodiversity hotspot) and rare forest biome, an unique wetland/estuarine lake system and the rugged mountain and coastal landscape provides diverse habitat in varied successional stages that consequently contribute to a reasonably diverse amphibian fauna. The GRNP also forms the eastern limit of a distinct zoogeographic zone, called the Cape Faunal Centre where the fauna gradually changes to include more Ethiopian elements characteristic of much of Africa. Regrettably, as with reptiles, knowledge of amphibians inhabiting the Park is limited.

Species Richness/Diversity

Systematic knowledge of the Western Cape Province’s (WCP) frogs is still in an alpha taxonomic stage with new species still being discovered and described (Turner et. al 2007). In the last five years in the WCP, at least three new species have been discovered (Turner et al. 2007) one species has been elevated from synonymy (Channing 2001) and another description is in preparation (Turner & Channing In prep.) There have also been two substantial revisions of the higher-level systematics of African frogs (Van der Meijden et al. 2005, Frost et al. 2006) and this has resulted in a number of name changes. This mirrors the situation in South Africa generally where current estimates put the total number of frog species at 115 (Minter et al ., 2004) that represents a notable increase (30%) from the last published total of 84 (Poynton, 1964).

The Garden Route National Park is considered to fall within one of the eight major centres of herpetofauna diversity in southern Africa (Branch, 1998) and forms part of the mountainous region stretching from the Cape in the south northwards to tropical Africa that is considered to function as a corridor along which various faunal taxa can move.

Formal amphibian checklists for the Park were derived from Branch & Hanekom (1987). These lists supplemented the preliminary checklist of Carruthers & Robinson's (1977). Amphibian checklists were derived from Von Breitenbach (1974), unpublished records from Cape Nature Conservation and Grindley (1985). Checklists were compiled by Branch and Hanekom (1987); Whitfield ( et al . 1983) and unpublished (SANParks) records of occurrence of species. All the frogs that might occur inside the Park are listed in Appendix C.

In total, the world contains more than 2500 species of frogs that are arranged in 270 genera and 33 families (Carruthers, 2001). South Africa has a total number of 115+ frog species (4.6 % of the world total), 33 genera (12.2 % of the world total) and 10 families (30 % of the world total). The WCP possesses fifty (43.5 % of SA) species, 19 genera and six families. The GRNP holds twenty-

management plan GRNP Appendices 32 two species (19.1% of SA and 44% of WCP), 10 genera and six families. Fourteen of the listed species (63.6%) have been formally recorded within the Park’s boundaries (Appendix A). The true species total is likely to be higher as the distribution ranges of number of species that could possibly occur within the Park end just short of Knysna.

Endemism

Eighteen species have varying degrees of endemism (Table 4). Nine species are Cape Floristic Kingdom endemics, namely Vandijkophrynus angusticeps , Capensibufo tradouwi , Heleophryne Regis , Afrixalus knysnae , Hyperolius horstockii , Breviceps fuscus , Breviceps montanus , Breviceps rosei and Strongylopus bonaespei . Five of these species are also endemic to the WCP and one is nearly endemic to the WCP. One species, Tomopterna delalandii , is a Cape Floristic Kingdom near-endemic. Two species are nearly-endemic to the ECP. Five species are endemic to South Africa and two species are southern Africa endemics. Afrixalus knysnae, a CFR endemic is entirely restricted to the southern Cape area (Branch and Hanekom 1987; Branch 1988; Minter et al . 2004). In addition, ten species (45.4%) are range restricted. Six species do not occur further east than Port Elizabeth and four species have western distribution limits, two of which do not occur west of George.

management plan GRNP Appendices 33

Table 4. Reptilian species endemic/near-endemic to the CFR, RSA and southern Africa.

Species C.F.R. endemic C.F.R. near- RSA Southern Africa endemic Afrixalus knysnae X Amietia fuscigula X Amietophrynus pardalis X X Amietophrynus rangeri X Breviceps fuscus X Breviceps montanus X Breviceps rosei X Cacosternum nanum X Capensibufo tradouwi X X Heleophryne regis

Hyperolius horstockii X X Semnodactylus wealii X Strongylopus bonaespei X Strongylopus fasciatus X Strongylopus grayii X

Tomopterna delalandii X Vandijkophrynus angusticeps Vandijkophrynus gariepensis

Conservation Status

Assessment of the conservation status of any fauna requires knowledge of its diversity, endemism, distribution, biology and habitat requirements, as well as information on threats that the animals face. Amphibian populations are declining throughout the world (Wake, 1991) due to a number of factors, including climate change, chemical pollution, habitat loss, and disease (Kiesecker et al. 2001; Blaustein et al. 2003). Furthermore, amphibians are considered to be very good indicators of pollution, acid rain and global warming (Branch, 1994; Channing, 1998; De Villiers, 1998) and it is thus very important that a verified list of all amphibians that occur inside the Park be compiled.

According to the Southern African Reptile Conservation Assessment (SARCA) that replaces the outdated SA Red Data List for Reptiles and Amphibians (Branch 1988) and uses modern IUCN criteria, only one species, namely Afrixalus knysnae , previously data deficient (RDB 2001) , is listed as Endangered. This species is declining rapidly and larger samples of advertisement calls are required for differences between similar species. The remaining 21 species are all in the least concern category (Table 10). However, of the least concern species, Vandijkophrynus angusticeps requires reassessment once taxonomic status of inland and montane poulations are determined

management plan GRNP Appendices 34

(Minter et al . 2004). Amietophrynus rangeri is declining and few recent observations have been made of Capensibufo tradouwi and surveys are recommended to confirm their current status. Hyperolius horstockii is protected against collection by provincial nature conservation regulations and several localities at which this species was recorded in the late 1970’s have disappeared under agriculture and urban development (e.g. Plettenberg Bay) and population monitoring in conservation areas is recommended.

Table 5. Conservation status of reptiles that occur in the GRNP

Order Family Scientific Name English SARDB IUCN CITE NEMBA Name Status Status S Category Vandijkophrynus Cape sand Least Anura Bufonidae angusticeps toad concern NL Null Vandijkophrynus Least Anura Bufonidae gariepensis Karoo toad concern NL Null Eastern Amietophrynus leopard Least Anura Bufonidae pardalis toad concern Null Null Amietophrynus Raucous Least Anura Bufonidae rangeri toad concern NL Null Null Tradouw Capensibufo mountain Least Anura Bufonidae tradouwi toad concern NL Null Null Heleophryne Southern Least Anura Heleophrynidae regis ghost frog concern NL Null Null Knysna Afrixalus leaf-folding Anura Hyperoliidae knysnae frog Endangered Endangered Null Endangered Hyperolius Arum lily Least Anura Hyperoliidae horstockii frog concern NL Null Null Hyperolius Painted Least Anura Hyperoliidae marmoratus reed frog concern NL Null Null Semnodactylus Rattling Least Anura Hyperoliidae wealii frog concern NL Null Null Plain rain Least Anura Brevicipitidae Breviceps fuscus frog concern NL Null Null Cape Breviceps mountain Least Anura Brevicipitidae montanus rain frog concern NL Null Null Rose's rain Least Anura Brevicipitidae Breviceps rosei frog concern NL Null Null Cacosternum Boettger's Least Anura Pyxicephalidae boettgeri caco concern NL Null Null Cacosternum Bronze Least Anura Pyxicephalidae nanum caco concern Null Null Common Least Anura Pipidae Xenopus laevis platanna concern NL Null Null Amietia Common Least Anura Pyxicephalidae angolensis river frog concern NL Null Null Anura Pyxicephalidae Amietia Cape river Least NL Null Null

management plan GRNP Appendices 35

fuscigula frog concern Strongylopus Banded Least Anura Pyxicephalidae bonaespei stream frog concern NL Null Null Strongylopus Striped Least Anura Pyxicephalidae fasciatus stream frog concern NL Null Null Strongylopus Clicking Least Anura Pyxicephalidae grayii stream frog concern NL Null Null Tomopterna Cape sand Least Anura Pyxicephalidae delalandii frog concern NL Null Null

Breviceps fuscus has a very restricted distribution and eastern limit Breviceps montanus is uncertain and Breviceps rosei’s habitat is continually being destroyed by alien plants. Montane populations of Xenopus laevis may be genetically distinct and Strongylopus bonaespei is intolerant of disturbance. Furthermore, although the bulk of the species are least concern, taxon data sheets were not completed for the majority of these species and these sheets need to be completed for each species. Although such a process would be unlikely to reveal any additional globally threatened species, it may reveal additional species that are threatened on a sub-regional or national scale.

One species namely, Afrixalus knysnae is listed as protected according to the NEMBA category (Table 2). No species are CITES listed but twenty species are also listed in the WCP ordinance schedule (Appendix I). A further nine frog species are categorised as conservation priority species in the WCP (Appendix I).

Ecological Studies

Ecological Knowledge of the amphibians of the GRNP is limited, consisting primarily of published and unpublished (SANParks) records of occurrence of species. The only papers produced on frogs in the GRNP till date was by Carruthers and Robinson (1977) and Branch and Hanekom (1987). Carruthers and Robinson (1977), provided information on the number and distribution of all species represented in the region. They found that the TNP included a wide variety of habitats suitable for amphibian life. A habitat of particular importance was found to be the grassy seepage areas adjacent to the sea where drainage from the slope collects here in thickly vegetated vleis before seeping through to the sea. Eleven frog species were identified that occupied ten habitat types. These habitat types were identified within four major topographical regions and the frog species associated with each of these was briefly described. Eight species were found to occupy the coastal strip, six were found in forest, ten on the plateau and four species were found to inhabit flood pans along the major rivers. Only three species occupied all four habitat types namely, Xenopus laevis , Strongylopus grayii and Amietia fuscigula . Two species were found in all habitats except major rivers, namely Cacosternum nanum and Semnodactylus wealii while Amietophrynus rangeri was found in all habitats except forest. Two species were only found on the coastal strip and plateau, namely, Strongylopus fasciatus and Hyperolius marmoratus . Three species were only found in one of the niches within the different habitats, e.g. Breviceps fuscus was entirely restricted to the forest floor and was the only species to inhabit this niche in the forest while Hyperolius horstockii and Afrixalus knysnae were only found in pans on the plateau.

management plan GRNP Appendices 36

Recent work (Minter, 2004) has shown that Breviceps fuscus also inhabits fynbos. Strongylopus grayii showed the widest distribution and was found in eight of the ten sub-habitats , and was the only species to inhabit the splash zone. Afrixalus knysnae (endangered) was only found in one locality, a shallow pan south of Covie village. Branch and Hanekom (1987) also recorded this species here only where juveniles were collected from restio stems along the edges of small pans. Advertisement calls of this species were recorded at Covie on 17 Sep 2002 (Braack and Burger unpubl. data). Pickersgill (1996; 2000) indicated that breeding occurs from October to December at Covie and Diepwalle and in late February in the De Vasselot section of the TNP. During the breeding season, males were seen calling from fallen branches and emergent vegetation just above the surface of small pools along the roadside and in forest clearings near Diepwalle (FitzSimons 1946). A Juvenile collected in early Oct at Covie and a female collected at Diepwalle in early December contained well developed ovarian eggs (Pickersgill, 2000). Tadpoles were also collected at Grootkop, Knysna. Of particular concern is that this species hs not been recorded from historical localities during atlas surveys in 2000 and 2001.

Branch and Hanekom (1987) produced an annotated checklist comprising 13 amphibia species and also briefly discuss their ecology and zoogeographic affinities. They found two additional species namely, Heleophryne regis and Hyperolius marmoratus . They indicated that many of the amphibian species occurring in the Park are representatives of the Cape temperate fauna (Broadly and Poynton, 1978) that are important future indicators of possible range contractions or expansions and that the Parks form important reserves for the conservation of these species.Since the TNP habitat types are found throughout he GRNP, it is reasonably expected that amphibian patterns are similar.

BIRDS

The GRNP with its combination of forest, fynbos, estuarine and marine habitats has a diverse avifauna. A total of 262 bird species have been recorded in the former WNP (Randall et al. 2007) and 257 in the former TNP (Skead & Liversidge 1957; Crawford 1983; N Hanekom & G McIlleron pers comm). These comprehensive park accounts, together with avifaunal studies in the former State Forests (e.g. Koen 1988), reveal that about 305 bird species have been recorded in what now comprises the Garden NP.

Included in this list are several species of special concern on account of their listing in the Red Data Book (cf Barnes 2000). These include 16 Endangered/Vulnerable species and a further 21 Near-threatened species of which 4 and 7 respectively are known to breed in GRNP.(Randall et al . 2007) The species of special concern include representatives of most of the major habitat types in the GRNP: marine (Cape cormorant Phalacrocorax capensis , crowned cormorant P.coronatus & African black oystercatcher Haematopus moquini ), wetlands (African marsh harrier Circus ranivorus, African finfoot Podica senegalensis, halfcollared kingfisher Alcedo semitorquata & African grass-owl Tyto capensis ), forest (Knysna warbler Bradypterus sylvaticus , Knysna

management plan GRNP Appendices 37 woodpecker Campethera notata & crowned eagle Stephanoaetus coronatus ) and general (peregrine falcon Falco peregrinus ).

The wetlands of the Touw and Swartvlei systems support one of South Africa’s most significant waterbird assemblages, which was the prime motivation for the listing of parts of the Touw system as a Ramsar wetland. About 110 bird species, over 35% of all species at GRNP, are dependent upon or primarily associated with wetlands. Rondevlei and Langvlei support the most diverse and abundant water bird communities (Boshoff & Piper 1992), with surveys on Langvlei indicating that at times water bird abundance can exceed 7000 individuals comprising 65 species (Boshoff & Palmer 1981). This abundance of water birds, and in particular Anatidae (ducks and geese) which on Langvlei alone at times exceeds 2000 individuals of nine species, represents the largest concentration of species and individuals along the southern and eastern Cape coasts (Underhill et al . 1980). Most water birds which occur on the lakes and estuaries have been observed to undergo short-term temporal, usually seasonal, variations in abundance (Boshoff et al. 1991a, 1991b, 1991c). Some of these variations can be related to changes in the systems, for example waterfowl abundance at the lakes has recently been shown to be determined by changes in the biomass of aquatic macrophytes (Russell et al. 2009).

The maintenance of water quality and quantity is fundamental to sustaining the ecological systems on which waterbird communities depend (Russell et al. 2009). Important criteria to maintain the terrestrial bird communities are features such as the conservation of sufficiently large tracts of land, where natural processes (e.g. fire) continue to function, and the existence of effective corridors to prevent situations where only islands of natural habitat remain.

MAMMALS

Introduction

The Garden Route National Park’s (GRNP) location within the fynbos (Floral Kingdom and Biodiversity hotspot) and rare forest biome, the inclusion of a marine and unique wetland/estuarine lake system and the rugged mountain and coastal landscape provides diverse habitat in varied successional stages that consequently contribute to a reasonably diverse mammal fauna. Moreover, the GRNP forms the eastern limit of a distinct zoogeographic zone, called the Cape Faunal Centre where the fauna gradually changes to include more Ethiopian elements characteristic of much of Africa. However, mammalian fauna biomass is low when compared to the summer rainfall regions mainly due to the fact that fynbos and forests in the southern Cape occur on nutrient-poor soils (Van Daalen 1980; 1981; 1984), and therefore have a low carrying capacity for sustaining large herbivores (Cody et al. 1983 in Rebelo 1992; Koen 1984). Additionally, several fauna species ranges, as with the flora species, end here which results in marginal environmental conditions that don’t favour high mammal densities. The Garden Route National Park is unique in the sense that it is largely unfenced and mammal species are able to freely move in and out of the Park.

management plan GRNP Appendices 38

Species Richness/Diversity

Mammal checklists have been produced over the years by Robinson (1976), Crawford (1982), Hanekom et al . (1987) and Pretorius et al . (1980) and Whitfield et al . (1983). Descriptions of the habits and distribution of 46 mammals recorded in the Knysna area is given by Von Breitenbach (1974) and Grindley (1985).

Eighty-seven species of mammal comprising fourteen orders and thirty-two families are known to occur within the Park of which eighteen species (21%) are restricted to the marine ecosystem. Sixty-nine species (79%) have been formally recorded (Appendix C). The total number of mammal species in South Africa is ± 290 (Lloyd, 2007), which means that only 30% of the species occurring in South Africa also occur in the GRNP. Similarly only 52% of the species occurring in the western Cape Province (WCP) occur in the GRNP. The majority, 52 (60%) species are small- bodied (less than 10 kg in mass). The largest groups are Rodentia with eighteen species, followed by Carnivores (16 species), Whippomorpha (14) and Chiroptera with 13 species that collectively account for 52% of the mammals occurring in the Park.

The true species total is likely to be higher as the distribution ranges of number of species that could possibly occur within the Park end just short of Knysna. For instance, Tatera afra (Cape gerbil), a western Cape endemic whose distribution range ends at Herold’s bay and Chrysochloris asiatica (Cape Golden Mole), a western Cape near-endemic whose range ends in Swellendam (Skinner and Chimimba, 2005); (Friedmann and Daly, 2004). Ichneumia albicauda (White-tailed mongoose), that reach their southern most limit in the eastern Cape at about 33 °27’S, have apparently been observed in the Wilderness National Park (Russel, 1998). This also applies to Leptailurus serval (Serval), a near-threatened species that has been reported from the Tsitsikamma State Forest. Stuart (1977, 1985) recorded that two servals were trapped and released in the far western Tsitsikamma National Park.

Another species Crocidura cyanea (Reddish-grey musk shrew), a data-deficient species that is shown to occur throughout the western and eastern Cape, Cistugo lesueuri (Lesueur's Wing-gland Bat) and Dasymys incomtus (Water Rat), both near-threatened, are other species that are likely to occur within the park boundaries. The same situation applies to marine mammals, the potential number of species even higher as many have distribution ranges that flank the park boundary while on their migratory routes.

Table 6. The number of species in each order is as follows:

ORDER FAMILIES SPECIES CR EN VU NT DD LC CITES AFROSORICIDA 1 3 0 0 0 1 1 1 0 CARNIVORA 6 16 0 1 0 1 1 12 8 CHIROPTERA 6 14 0 0 0 6 0 8 0 EULIPOTYPHLA 1 5 0 0 0 1 4 0 0 HYRACOIDEA 1 1 0 0 0 0 0 1 0

management plan GRNP Appendices 39

LAGOMORPHA 1 2 0 0 0 0 0 2 0 MACROSCELIDAE 1 2 0 0 0 0 0 2 0 PRIMATES 1 2 0 0 0 0 0 2 2 PROBOSCIDEA 1 1 0 0 0 0 0 1 1 RODENTIA 4 18 0 1 0 0 1 16 0 RUMINANTIA 1 7 0 0 1 0 0 6 1 SUIFORMES 1 1 0 0 0 0 0 1 0 TUBULIDENTATA 1 1 0 0 0 0 0 1 0 WHIPPOMORPHA 6 14 0 0 3 1 5 5 13 32 87 0 2 4 10 12 58 25

Several species that historically would have occurred in the Garden Route ecosystem are now locally extinct – Panthera leo (Lion), Diceros bicornis (Black rhinoceros), Hippopotamus amphibius (Hippopotamus), Syncerus caffer (Buffalo), Tuarotragus oryx (Eland), Alcelaphus buselaphus (Red hartebeest) and Equus zebra zebra (Cape Mountain Zebra) (Lloyd 2002; Skead, 1980) The last small relict population of Knysna elephants ( Loxodonta africana ) face a similar outcome despite conservation intervention.

Endemism

Although not in quite the same proportion as the rest of South Africa, the GRNP also has endemic mammals and even though the Park’s forest and fynbos environments do not exhibit high levels of mammalian biodiversity (in contrast with many forest habitats elsewhere in the country) there are a few mammalian species and subspecies that are nearly endemic to these forests. Philantomba monticola (blue duiker) and Potamochoerus porcus (bushpig) are species whose major strongholds are within the forests but both species are also found in adjacent thicket or coastal scrub environments.

Twelve species, three of which are Near-Threatened, have varying degrees of endemism (Table 7). Two of these species are endemic to the western Cape and the Cape Floristic Kingdom - Acomys subspinosus (Cape spiny mouse) and Bathyergus suillus (Cape Dune Molerat). Four species are endemic to the Cape Floristic Kingdom and near-endemic to the western Cape - Myosorex longicaudatus (Verreaux’s mouse), Chlorotalpa duthieae (Duthie's golden mole), Myomyscus verreauxi (Verreaux's mouse) and Amblysomus corriae (Fynbos Golden Mole). Chlorotalpa duthieae is only found along the southern Cape coast from George to Humansdorp. One species is both near-endemic to the Cape Floristic Kingdom and western Cape - Raphicerus melanotis (Grysbok). The five remaining species are endemic to South Africa. One small mammal that is largely confined to this ecoregion is the long-tailed forest shrew ( Myosorex longicaudatus , NT) (Hilton-Taylor 2000).

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Table 7. Mammalian species that are endemic/near-endemic to the Western Cape Province (W.C.P), Cape Floristic Kingdom (C.F.K.) and South Africa (SA).

Species C.F.K. W.C.P. (NE) SA W.C.P .

Acomys subspinosus X X Bathyergus suillus X X Myosorex longicaudatus X X Chlorotalpa duthieae X X Myomyscus verreauxi X X Raphicerus melanotis X (ne) X Amblysomus corriae X X Amblysomus hottentotus X Elephantulus edwardii X X Georychus capensis X Graphiurus ocularis X Rhinolophus capensis

Conservation Status

According to the new IUCN categories (IUCN 2001), that were used to evaluate the threat status of the South African mammals (Friedmann and Daly 2004), the Park protects populations of twenty- eight red data book species that originate from seven orders and thirteen families (Table 8 and Figure 2 and 3). In the threatened categories, 7.1% of the Parks mammals are listed as endangered (EN) and 14% as vulnerable (VU).

Apart from the Parks threatened species (CR, EN, and VU), 36 % and 42% of the Parks mammals are listed in the near threatened (NT) and data deficient (DD) categories respectively. Effectively this means that 34 % of the Parks mammals either are, or could potentially soon be, facing a threat to their future survival (Lloyd 2002). Only 58 (66%) mammals fall in the Least Concern (LC) category and one species, Lobodon carcinophagus (Crabeater seal), has not yet been properly evaluated. Whippomorpha (9), Chiroptera (6), Eulipotyphla (5) and Carnivora (3) account for the bulk (82%) of the red listed species. Only four orders - Whippomorpha, Carnivora, Rodentia and Ruminantia are in the threatened category (Threatened and Vulnerable). Twenty-five species are CITES listed (Table 7 and 8), 76% of which fall in the Appendix II category. Whippomorpha represents the bulk of the CITES listed species with thirteen (52%) species, one of which is endangered and four vulnerable. Eight (32%) Carnivora species are CITES listed, one of which is the near threatened Honey badger, which is also the Parks only Appendix III listed mammal species (Table 8).

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Figure 2. Conservation status of GRNP mammals EN = Endangered, VU = Vulnerable, NT = Near Threatened and DD = Data Deficient

14

12 10 8

6

4

2

Number of species 0 EN VU NT DD IUCN CATEGORIES

Figure 3. Conservation status of GRNP mammals EN = Endangered, VU = Vulnerable, NT = Near Threatened and DD = Data Deficient

10

9

8

7

6

5 IUCN LISTINGS 4 3 2

1

0 AFROSORICIDA CARNIVORA CHIROPTERA EULIPOTYPHLA RODENTIA RUMINANTIA WHIPPOMORPHA

Order

Noteworthy is that Whippomorpha (marine mammals) make up 50% of the threatened species category and the remaining red listed species fall in the data deficient category, most probably meaning that this statistic is most likely and under-estimation. In addition, Mirounga leonina (Southern elephant seal) an endangered species is marine dependent. Of further concern is Chiroptera, where 43% of red listed species in this order are near threatened.

Table 8. Red listed mammals that occur in the GRNP

Order Family Scientific name Common name IUCN Status

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Ruminantia Bovidae Philantomba monticola Blue Duiker VU Whippomorpha Balaenidae Balaenoptera edeni Bryde's whale VU Whippomorpha Physeteridae Physeter catodon Sperm whale VU Whippomorpha Delphinidae Sousa chinensis Humpback dolphin VU Afrosoricida Chrysochloridae Amblysomus corriae Fynbos Golden Mole NT Carnivora Mustelidae Mellivora capensis Honey Badger NT Chiroptera Vespertilionidae Kerivoula lanosa Lesser woolly bat NT Chiroptera Vespertilionidae Miniopterus fraterculus Lesser long-fingered bat NT Chiroptera Vespertilionidae Miniopterus schreibersii Schreibers' long-fingered bat NT Chiroptera Vespertilionidae Myotis tricolor Temminck's hairy bat NT Chiroptera Rhinolophidae Rhinolophus capensis Cape horseshoe bat NT Chiroptera Rhinolophidae Rhinolophus clivosus Geoffroy's horseshoe bat NT Eulipotyphla Soricidae Myosorex longicaudatus Long-tailed forest shrew NT Whippomorpha Balaenopteridae Megaptera novaeangliae Humpback whale NT Carnivora Phocidae Mirounga leonina Southern elephant seal EN Rodentia Muridae Mystromys albicaudatus White-tailed rat EN Afrosoricida Chrysochloridae Amblysomus hottentotus Hottentot golden mole DD Carnivora Mustelidae Poecilogale albinucha African striped weasel DD Eulipotyphla Soricidae Crocidura flavescens Greater red musk shrew DD Eulipotyphla Soricidae Myosorex cafer Dark-footed forest shrew DD Eulipotyphla Soricidae Myosorex varius Forest shrew DD Eulipotyphla Soricidae Suncus infinitesimus Least dwarf shrew DD Rodentia Muridae Grammomys dolichurus Woodland thicket rat DD Whippomorpha Ziphiidae Mesoplodon densirostris Blainville's beaked whale DD Whippomorpha Ziphiidae Mesoplodon hectori Hector's beaked whale DD Whippomorpha Delphinidae Orcinus orca Killer whale DD Whippomorpha Delphinidae Tursiops truncates Bottlenose dolphin DD Whippomorpha Delphinidae Grampus griseus Risso's dolphin DD

The two species considered endangered are Mirounga leonine (Southern elephant seal) and Mystromys albicaudatus (White-tailed rat), the former a CITES listed Appendix IIA species. The four mammalian species considered Vulnerable (V), are Balaenoptera edeni (Bryde's whale), Philantomba monticola (Blue Duiker), Physeter catodon (Sperm whale) and Sousa chinensis (Humpback dolphin), all CITES listed (Table 4).

The ten Near Threatened (NT) species are Amblysomus corriae (Fynbos golden mole), Kerivoula lanosa (Lesser woolly bat), Megaptera novaeangliae (Humpback whale), Mellivora capensis (Honey Badger), Miniopterus fraterculus (Lesser long-fingered bat), Miniopterus schreibersi (Schreibers' long-fingered bat), Myosorex longicaudatus (Long-tailed forest shrew), Myotis tricolor (Temminck's hairy bat), Rhinolophus capensis (Cape horseshoe bat) and Rhinolophus clivosus (Geoffroy’s Horseshoe Bat). Two of these species, namely Megaptera novaeangliae and Mellivora capensis are CITES listed. Two more species, Amblysomus corriae and Myosorex

management plan GRNP Appendices 43 longicaudatus are endemic to both the western Cape Province (WCP) and the Cape Floristic Kingdom (CFK). The third species, namely Rhinolophus capensis is endemic to South Africa (Table 9).

Table 9. CITES listed mammals that occur in the GRNP.

Scientific name Common name CITES 1986 Aonyx capensis African clawless otter Appendix II Arctocephalus pusillus South African Fur Seal Appendix II Arctocephalus tropicalis Subantarctica Fur Seal Appendix II Balaenoptera edeni Bryde's whale Appendix I Caracal caracal Caracal Appendix II Cercopithecus pygerythrus Vervet Monkey Appendix II Delphinus delphis Short-beaked common Dolphin Appendix II Eubalaena australis Southern right whale Appendix I Felis silvestris African wild cat Appendix II Kogia breviceps Pygmy sperm whale Appendix II Kogia sima Dwarf sperm whale Appendix II Loxodonta africana African Elephant Appendix II Megaptera novaeangliae Humpback whale Appendix I Mellivora capensis Honey Badger Appendix III Mesoplodon densirostris Blainville's beaked whale Appendix II Mesoplodon hectori Hector's beaked whale Appendix II Mirounga leonina Southern elephant seal Appendix II Orcinus orca Killer whale Appendix II Panthera pardus Leopard Appendix I Papio hamadryas Chacma baboon Appendix II Philantomba monticola Blue Duiker Appendix II Physeter catodon Sperm whale Appendix I Sousa chinensis Humpback dolphin Appendix II Stenella coeruleoalba Striped dolphin Appendix II Tursiops truncates Bottlenose dolphin Appendix II

Apart from the threatened species (EN, and VU), a further twelve of the Park’s mammal species have been placed in the Data Deficient (DD) category, namely Amblysomus hottentotus (Hottentot golden mole), Crocidura flavescens (Greater red musk shrew), Grammomys dolichurus (Woodland thicket rat), Grampus griseus (Risso's dolphin) Mesoplodon densirostris (Blainville's beaked whale), Mesoplodon hectori (Hector's beaked whale), Myosorex cafer (Dark-footed forest shrew), Myosorex varius (Forest shrew), Orcinus orca (Killer whale), Poecilogale albinucha (African striped weasel), Suncus infinitesimus (Least dwarf shrew) and Tursiops truncates (Bottlenose dolphin).

Ecological Studies

management plan GRNP Appendices 44

Adequate information remains a prerequisite for sound management (Seydack, 1991) and fundamental to the effective management and conservation of any species, is a reliable estimation of its population size, distribution and trends. However, population sizes and conservation status of most of the important species are largely unknown. Of the specialized studies prepared on mammals in GRNP, much emphasis has been on the large forest mammals.

Table 10: Mammals of the GRNP

ORDER: RUMINANTIA One family (Bovidae) and seven species are found in this order namely, Oreotragus oreotragus (Klipspringer), Pelea capreolus (Grey rhebok), Raphicerus campestris (Steenbok), Raphicerus melanotis (Cape grysbok) and Sylvicapra grimmia (Common duiker) are considered prominent fynbos species, while Philantomba monticola (Blue duiker) and Tragelaphus scriptus (Bushbuck) are predominantly forest species. Studies have only been completed for the forest- restricted species namely Blue Duiker and Bushbuck. The fynbos species are very rarely seen and their ecology should be investigated. Family: Bovidae Philantomba monticola (Blue Duiker) The Blue duiker reaches its most southerly distribution limit in the Afromontane forests of the southern Cape (Crawford, 1984). This species is listed as the only red data book (Vulnerable), National Environmental Management: Biodiversity Act, 2004 (Vulnerable) and CITES (Appendix II) species in the order due to population declines as a result of habitat fragmentation and unsustainable utilisation. The species has fortunately received some attention and particularly well studied were population densities after a noticeable widespread population crash from about 1960 demanded an explanation.

A pellet group count in the Knysna forests for bushbuck and blue duiker was first carried out under von Gadow (1978) and the applicable indices for defecation, and decay rates determined. He estimated a population density of between 0.09 and 0.55 duikers per hectare, found that blue duiker are more numerous in the larger, continuous forests and detected mutual avoidance between blue duiker and bushbuck. In addition, he found that blue duiker avoid artificially slashed onderbos areas, significantly favour undecomposed litter and the higher altitude forests.

A pilot project employing auto-triggering camera units was tested as a method of studying larger mammals in the Knysna Forests (Seydack 1984). Of 596 photographic animal records, only fifteen photographic records (2.5%) were obtained for this species that was not enough to estimate densities. In 16 years, a mere 17 (3%) of 575 mammal observations made in the Knysna and Tsitsikamma forests were of blue duiker.

Crawford (1984) investigated activity, group structure and lambing of blue duiker in the Tsitsikamma National Park forests and found that they were active throughout the day and spend

management plan GRNP Appendices 45 about eight hours a day foraging. Crawford (1984) implicated predation as the reason for decline of blue duiker densities. Seydack (1977) also reported noticeable increases in caracal sightings in the Goudveld area and a decline in blue duiker sightings.

Hanekom and Wilson (1991) determined blue duiker densities in the Tsitsikamma National Park forests and probable factors limiting these populations. They found that the population density estimates of the blue duiker in the park were similar to those determined from four southern Cape forests and two Zairian forests but had at least three times lower population densities than that recorded by Bowland (1990); Bowland & Perrin (1994) in KwaZulu - Natal. They also found that limiting and unpredictable resource availability, for instance mast fruiting reproductive strategies by trees (Janzen 1974) were most probably responsible for the low duiker population densities. Moreover, predator scats were analysed and only 26 leopard scat piles were found during a two-year study period that revealed blue duiker remains in only 27% of leopard ( Panthera pardus ) and 7% of caracal (Caracal caracal) scats.

Hanekom and Wilson (1991) reported that in 95% of feeding observations, duikers foraged on the ground and often in areas where ground cover was less than 10%. In addition, they also found that dicotyledonous leaves represent about 57% of analysed stomach contents and that fruits and leaves comprised about 71% (by mass) of ingested food. Canthium obovatum (Kwar) was the most important species selected and the fruits of Dietes iridioides and Canthium mundianum were also taken and, together, fruits and leaves comprised about 71% of ingested food.

Seydack et al . (1998) showed that forest ungulate population fluctuations were driven by naturally varying climatic trends which has an indirect effect on metabolism by altering Nitrogen/Carbon ratios of dietary items, favouring bushbuck populations in warmer climates and vice versa.

Skinner and Chimimba (2005) point out that in blue duiker specimens from the southern coastal forests are unique in that the body is dark blue-grey and the limbs light rusty-brown.

management plan GRNP Appendices 46

Tragelaphus scriptus (Bushbuck) Bushbuck, which are one of the most common of the larger forest mammals, are true forest dwellers (Odendaal 1977, Seydack, 1984).

Odendaal & Bigalke (1979) determined the home range size in the Knysna forest, where the density of bushbuck was less than 5 individuals/km 2, which is approximately one-quarter the density of bushbuck from the savanna biome. The mean homerange size was 102 hectares (range=14.6-174.3), and the smallest (14.6ha) was that of a lactating female and could probably be attributed to the concealment behaviour of bushbuck young.

Seydack (1984) found similar densities recorded 187 (32%) photographic records in a total of 596 photographic records of 14 different fauna species, from which he determined a population density of 5 and 7 animals/km2.

Odendaal & Bigalke (1979) investigated social organisation of the species, of 776 animals observed in the Knysna forest, 58% occurred singly, 32% in pairs and only 10% in groups of three or more. The authors also discovered that single males, single females, and male/female associations accounted for 71% of all their observations. Since these associations were also detected in Zimbabwe (Wilson & Child, 1964) and northern Botswana (79% - Elder & Elder, 1970), it was postulate that the three associations play an important role in the social organisation of the species (Odendaal & Bigalke 1979).

Odendaal & Bigalke (1979) suggested that a negative correlation exists between the size of bushbuck home ranges and population densities. They also reported that home ranges of adult males overlap considerably, as do the home ranges of males and females.

Odendaal (1983) analysed the stomach contents of 25 animals in the Knysna Forest and recorded a list of 47 food plants. Browse comprised 83% of the diet while monocotyledons comprised only 10%. Odendaal (1983) suggested that mycophagy might be essential to guarantee nutritional adequacy in the diet.

Odendaal & Bigalke (1979) found that although lambing peaks in April, August and November, the fact that the male-female associations are formed throughout the year ensures continual lambing.

As with blue duiker, southern Cape bushbuck (spp. sylvaticus ) are morphologically unique from the northern population in having much darker, almost black coat hair.

No local knowledge is available on the remaining five mammal species in this order, one of which (Cape grysbok) is near endemic to the western and eastern Cape and the Cape Floristic Kingdom.

management plan GRNP Appendices 47

ORDER: CARNIVORA This order is second in terms of species diversity with six families and 16 species. namely: Mustelidae - Aonyx capensis (African clawless otter), Mellivora capensis (Honey Badger), Ictonyx striatus (Striped Polecat) and Poecilogale albinucha (African striped weasel); Otariidae - Arctocephalus pusillus ( South African Fur Seal) and Arctocephalus tropicalis (Subantarctica Fur Seal); Felidae – Panthera pardus (leopard) and Caracal caracal (caracal) and Felis silvestris (African wild cat); Herpestidae - Atilax paludinosus (Marsh mongoose) and Herpestes ichneumon (Large grey mongoose); Viverridae - Galerella pulverulenta (Cape grey mongoose), Genetta genetta (Small-spotted genet) and Genetta tigrina (Large-spotted genet); Phocidae - Lobodon carcinophaga (Crab-eating seal) and Mirounga leonine (Southern elephant seal). Family: Mustelidae Aonyx capensis (African clawless otter) There are only thirteen species of otter’s worldwide and most of them are threatened. Otters are excellent indicators for wetlands, because they are top predators in various semi-aquatic habitats. In Africa four species of otters occur, but we know very little about them (IUCN/SSC). The GRNP, with its diverse aquatic habitat, provides outstanding habitat for this species. Apart from knowledge accumulated from formal ecological studies examined below, monitoring programmes in the Park have also contributed to knowledge. For instance, reported sightings from all over the GRNP has aided in determining the range and evidence of breeding has been reported in Lelievlei nature reserve. In 16 years, 10 (1.7%) of 575 mammal observations made in the Knysna and Tsitsikamma forests were of otter, 70 % of which were in marine habitat. Prey remains in scats indicate a high incidence of crab.

The Tsitsikamma Coastal National Park (TCNP) has been the main focus of studies on this habitat specific, charismatic species that is CITES (Appendix II) listed due to persecution, hunting and accidental trapping. These studies concentrated on the marine habitat only. Van der Zee (1982); Arden-Clarke (1983) reported that approximately thirty ( Aonyx capensis ) Cape clawless otters occur along the coast of the TCNP. Van der Zee (1979,1981a) reported ground densities of 1 otter/2km of coastline that appear higher than densities at freshwater habitats at 1otter/8-10km of river bank (Butler and du Toit, 1994). Arden-Clarke (1983) found that the main periods of activity were from 20h00 to 22h00. The overall recorded foraging ranges were between 8.5 and 19.5 km (Arden-Clarke 1983). Home ranges of males and females overlap (van der Zee 1982; Arden-Clarke 1986) and temporal instead of spatial avoidance may occur (van der Zee 1982). Nightly movements may reach 13km (Taylor, 1970), and males are believed to move longer distances than females.

Van der Zee (1979, 1981) analysed over 1 100 spraints and recorded over 30 species of crabs, fish and octopus. Approximately 80 % of prey items consisted of four species, namely Plagusia chabrus (Cape rock crab) that had the highest frequency of 27.9%, followed by Chorisochismus dentex (Rocksucker fish) at 16.5% and Cyclograpsus punctatus (brown rock crab) at 10.4%. The fourth species, Octopus granulatus (octopus) had a frequency of 7.4%. Clinus cottoide s (small

management plan GRNP Appendices 48 rockfish) had a frequency of 4.4% and the remaining food items all had a relative frequency of less than 2.5%. Arden-Clarke 1983) reported octopus as the third (15.1%) most important prey item. Spraints occur mostly on rocks and boulders and <50 m from a freshwater source (van Niekerk et al. 1998).

Van der Zee (1982) reported that most observations (78%, n = 124) were of single otters, whereas groups of 2, 3, and 4 comprised 12%, 9%, and 1% of observations, respectively. On occasion, groups of males may forage together in a clan-type organization (Arden-Clarke 1986). Of 56 observations made by Arden-Clarke (1986) in the TCNP, the mean group size was 1.71. Thirty-five of these observations were of solitary animals, five of pairs, eight of three, three of four and one of a group of five.

Dens of Cape clawless otters were typically near an abundant food supply, near bushy areas, near (<15 m) fresh water, and never located >50 m from shore (van der Zee 1982; van Niekerk et al. 1998). Average use of all den sites (n =65) was 32%, and each site was visited on average every 3 days (van der Zee 1982). Distance between den sites averaged 1.0 km, but varied from 0.3 to 2.4 km and each site was visited on average every three days (van der Zee 1982). Arden- Clarke (1986) traced 15 couches, used by the same individual, in a 12.8km stretch of coastline and reported a density of 2.1 dens/km of coastline and that several Cape clawless otters may share dens.

The most important factor threatening Cape clawless otters is increasing human populations and associated habitat changes (Butler 1994; Rowe-Rowe 1986, 1990, 1995; Rowe-Rowe et al. 1995; Van der Zee 1982). Mellivora capensis (Honey Badger) Of 596 photographic records obtained during a census project in the Knysna forests, 120 (20%) were of this species. Of the 120 records, 33% were during daylight, 13% at dawn or dusk and 54% at night (Seydack, 1984). This omnivorous animal has a wide food spectrum, but is particularly fond of honey and bee larvae. Of 575 mammal observations reported in the Knysna and Tsitsikamma forests over 16 years, 38 (7%) were of this species. Since this species is of conservation significance (Table 2 and 3) ecological studies are overdue. Seydack (1991) stressed that the conservation of this species entails maintenance of suitable habitat such as indigenous forests and that further studies on this animal, particularly within the forest environment, would be highly desirable. Family: Viverridae Galerella pulverulenta (Cape grey mongoose) This species is widespread in the forested and fynbos areas (Robinson, 1976; Crawford et al. 1983). They are predominantly solitary but can be found in groups of up to five (Stuart 1981,1991). Crawford et al. (1983) made some observations on this species in the Tsitsikamma National Park and reported that the species is common in dry scrub forests, uncommon in dry high forests and possibly absent from moist and wet high forests. Eighty nine percent of sightings in the Tsitsikamma National Park were of solitary animals, ten percent are of pairs and

management plan GRNP Appendices 49 one percent are of groups of three (Crawford et al. 1983). They also found that a single animal ranged over 75ha and that the species was most active during early morning and late afternoon and seasonally from late autumn through to early spring, with adults tending to remain in a given territory. In addition, they reported that sociality is highest during summer and pairs are also seen in July and August. This species was present in prey species collected at Crowned eagle nests in the GRNP. Genetta genetta (Small-spotted genet) Seydack (1977) indicates that this species is abundant at Goudveld (5 individuals caught in 16 trapnights) and occurs in fynbos, forest and plantation. In addition (Seydack 1984) recorded 84 (14%) photographs out of a total of 596 in a photographic census of forest mammals. Of 575 mammal observations reported in the Knysna and Tsitsikamma forests over 16 years, 27 (5%) were of this species. Family: Felidae Felis silvestris ( African wild cat) This cat’s presence was detected by Pretorius et al . (1980) and although no local knowledge is available for this species, it is mentioned here since it is very rarely seen and its conservation status needs to be established. Panthera pardus (Leopard ) The leopard, which is listed as vulnerable in the National Environmental Management: Biodiversity Act, 2004 occurs widely, but sparsely, throughout southern Africa and in appreciable numbers only in conservation areas or remote, inaccessible mountains and forests (Von Richter, 1972) and is essentially a forest animal (Kingdon, 1977) but can tolerate a wide variety of habitats.

The ecology of the southern Cape leopard has not been investigated in any detail and their population status remains largely unknown. . Research on the Cape leopard in Stellenbosch and the Cedarberg indicate that this species occupies larger home-ranges in low resource environments. Hanekom and Wilson (1991) investigated the diet of Tsitsikamma leopards by analysing 26 scat piles. The following proportions of prey remains were discovered – birds and blue duiker (27%), rodents and dassie (15%), insects and bushpig (12%), bushbuck, baboon, porcupine (8%) and caracal (4%).

Of 596 photographic records obtained during a census project in the Knysna forests, 120 (20%) photos were of this species. Of the 120 records, 33% were during daylight, 13% at dawn or dusk and 54% at night (Seydack, 1984). Of 596 photographic records obtained during a census project in the Knysna forests, 6 (1%) recordings were made representing three different individuals, two adults and one sub-adult (Seydack, 1984). In 16 years, 32 (6%) of 575 mammal observations made in the Knysna and Tsitsikamma forests were of leopard (Herd, Unpubl records). Of these sightings, quite a few have been of young animals, and of substance was a sighting of a female with four cubs in the Fisantehoek forest.

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Myers (1976) advised that in southern Africa viable leopard populations would be confined to conservation areas by the turn of the century because of rapid habitat destruction. The GRNP protects a large part of this leopard refuge. More area-specific quantitative information is needed on this species for rational conservation management Seydack (1991). This is particularly so as conservation areas are not continuous and leopard ranges often overlap with “hostile” agricultural areas (Seydack, 1991).

No knowledge is available on the remaining nine mammals in this order. Of these, three are CITES listed (Appendix II) and African striped weasel is data deficient. Caracal caracal (Caracal) Sightings of caracal occur in most habitats throughout the park. It is said that caracal were absent from the area and have moved in from the drier northern areas after drought in the Karoo (Seydack, 1977). The creation of networks of roads over the years have no doubt aided in the distribution of caracal deeper into forest for instance. Seydack (1977) also reports a noticeable increase in caracal sightings in the Goudveld area and a decline in blue duiker sightings. Consequently, caracal was also considered as one of the reasons for blue duiker population declines (Liversidge, 1966; Robinson, 1976; Crawford and Robinson, 1984).

Seydack (1984) recorded nine photographs of the species between December 1978 and November 1981 in the Goudveld forest. Twenty sightings were made in the western sector of the Tsitsikamma National Park between May 2000 and September 2003 (A.R. Unpubl records). In the last 16 years, 46 (8%) out of 575 mammal observations made in the Knysna and Tsitsikamma forests were of caracal (Unpubl records). Hanekom and Wilson (1991) investigated diet by analysing twelve scat piles. The following proportions of prey remains were discovered - Rodents (43%), birds and insects (29%), dassie (14%), blue duiker (7%) and bushpig (7%). Family: Phocidae Mirounga leonina (Southern elephant seal) Seal have been recorded along the beaches of the Wilderness and Tsitsikamma National Parks though none are permanently resident in the area. The majority of beached individuals are juvenile or sub-adult Cape fur seals, which are typically in an emaciated condition. In 1992 a single adult male Southern elephant seal (Endangered) hauled out at Gericke’s Punt, where over a period of five weeks it underwent its annual moult (Randall 1992). ORDER: AFROSORICIDA This order comprises one family and three species namely, Amblysomus corriae (Fynbos Golden Mole), Amblysomus hottentotus (Hottentot golden mole) and Chlorotalpa duthieae (Duthie's golden mole).

Family: Chrysochloridae Amblysomus corriae (Fynbos Golden Mole)

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The Fynbos Golden Mole, declared near-threatened and endemic is of considerable conservation importance (Friedmann and Daly, 2004). Survey studies, taxonomic research and life history aspects are urgently required nationally for this species. It is essential to determine the current status of the species and the contribution that the Park can make to its conservation. ORDER: CHIROPTERA This order comprises six families and fourteen species. Emballonuridae - Taphozous mauritianus (Mauritian Tomb bat); Molossidae - Tadarida aegyptiaca (Egyptian Free-tailed Bat); Nycteridae - Nycteris thebaica (Egyptian slit-faced bat); Pteropodidae - Rousettus aegyptiacus (Egyptian rousette); Rhinolophidae - Rhinolophus capensis (Cape horseshoe bat) and Rhinolophus clivosus (Geoffroy's horseshoe bat); Vespertilionidae - Kerivoula lanosa (Lesser woolly bat), Miniopterus fraterculus (Lesser long-fingered bat), Miniopterus schreibersii (Schreibers' long-fingered bat), Myotis tricolor (Temminck's hairy bat) Neoromicia capensis (Cape serotine bat) and Pipistrellus hesperidus (African pipistrelle). Family: Nycteridae Substantial numbers of Egyptian fruit bats Rousettus aegyptiacus were recorded in a cave in the Storms River gorge (ca. 3 000 individuals) and Elands River gorge (ca. 300 – 400) (Herzig- Straschel & Robinson 1978, Swart 2002). This species feeds on the fruit of plants such as essenhout, saffron, poison bush and mistletoe, and their nocturnal activity is significantly longer in summer than in winter. Parturition apparently occurs between October and June, with a peak in December (Herzig- Straschel & Robinson 1978). A number of insectivorous bats are also present and it is likely that at least some of the species are highly dependent upon the forest for roosting and hunting habitat. Numerous river valleys, coastal caves, cliffs and historic mineshafts in the Millwood area provide exceptional bat roosting habitat. Four species, Rousettus aegyptiacus, Rhinolophus clivosus (near threatened), Rhinolophus capensis and Miniopterus schreibersii were confirmed for the Farleigh area (Seydack, 1977; Lück, 2000). Caves along the Harkerville and Tsitsikamma coastlines harbour numerous bat populations that are currently being researched by UCT. Of conservation importance are the six near threatened species, one of which is endemic. Additional surveys are urgently needed. ORDER: EULIPOTYPHLA The five species in this order, all of which belong to the family Soricidae, namely Crocidura flavescens (Greater red musk shrew), Myosorex cafer (Dark-footed forest shrew), Myosorex longicaudatus (Long-tailed forest shrew), Myosorex varius, (Forest shrew) and Suncus infinitesimus (Least dwarf shrew) are all of conservation importance. One species is near threatened and near endemic (Long-tailed forest shrew) and the remaining bat species are data deficient (Friedmann and Daly, 2004). There is however, little knowledge on the status and ecology of shrews in the Park. Myosorex longicaudatus is a recently discovered species that is known from only five localities in the southern parts of the western Cape (Cameron, 1982; Skinner and Chimimba, 2005), two of the sites being the Lottering and Diepwalle Forests. Specimens from Lottering were found to be more reddish-brown in colour (Dippenaar, 1995). It is essentially a forest animal (specialist) requiring moist habitat with ferns. Interestingly, a sub- species ( Myosorex boosmani ) found only in the Boosmansbos wilderness area is separated from

management plan GRNP Appendices 52 the Knysna population by the dry Gouritz river valley. Survey studies and behavioural research need investigation (Friedmann and Daly, 2004). ORDER: HYRACOIDEA One family, Procaviidae and one species, Procavia capensis only represent this order. The species is restricted to rocky habitat in the Park, particularly along the coast. Fairall and Hanekom (1987) monitored yearly fluctuations in the rock hyrax Procavia capensis in the Storms river restcamp area. The numbers increased steadily in late 1970s and early 1980s, before stabilizing in the mid 1980s. A simple simulation model was constructed and applied to evaluate different management options (Fairall and Hanekom 1987). Rock hyrax (Procavia capensis ) contributed to 25 percent of the prey individuals recorded from six crowned eagle nests in the southern Cape forest biome (Boshoff et al. 1994). ORDER: LAGOMORPHA One family and two species represent the order. Lepus capensis (Cape hare) and Lepus saxatilis (Scrub hare) have been recorded at Goudveld and Wilderness but are exceptionally scarce throughout the park and need to be surveyed. ORDER: MACROSCELIDAE One family (Macroscelididae) and two species, one a South African endemic represent the order. Elephantulus edwardii (Cape Rock Elephant-shrew) and Macroscelides proboscideus (Round- eared Elephant-shrew) are still to be recorded in the park and need to be surveyed. ORDER: PRIMATES One family (Cercopithecidae) and two species, namely Cercopithecus pygerythrus (Vervet Monkey) and Papio hamadryas (Chacma baboon) that are both CITES Appendix II listed, represent the order. ORDER: PROBOSCIDEA Represented by one family (Elephantidae) and species ( Loxodonta africana ), the rapidly diminishing elephant population has been the best documented (see Phillips (1925), Wildlife Society (1970), Carter (1970,1971), Thesen (1981), Oliver (1982) and MacKay (1983). These elephants had a much wider range before hunting, agriculture and industrialisation confined them to the forests. By 1994 it was thought that four elephants still roamed the forests but an intensive search confirmed the presence of just one adult cow. An unsuccessful attempt was made in July 1994 to increase the relict population by introducing three orphaned female juveniles from Kruger National Park.

Numerous studies have been undertaken on the ecology of the Knysna elephants. Phillips (1925) describes their history and habits. Von Gadow (1973) made observations on the utilization of indigenous trees by the Knysna elephants. Koen (1983) performed a study of the distribution, population composition, movements and feeding of the Knysna elephants. Koen (1983) also investigated seed dispersal by the Knysna elephants. Seydack (1983) explored aspects of elephant habitat suitability in the Knysna forest and surroundings. Koen (1988) investigated

management plan GRNP Appendices 53 trace elements and some other nutrients in the diet of the Knysna elephants. Koen ( et al. 1988) reported on macronutrients in plants available to the Knysna, Addo, and Kruger National Park elephants. Milewski (2002) reported on the diet of the Knysna and introduced Kruger elephant’s and provided evidence that elephants routinely eat the nutrient-poor, heathy shrubs and wiry grasslike plants of fynbos. Milewski (2002) indicated that seeds of exotic species seldom germinated from their faeces and suggested that elephants may be useful in controlling invasive exotic trees in the southwestern Cape. He also established that the last native elephants ate eleven out of approximately 35 species of forest trees and shrubs available in their home range.

The critical question is why the Knysna elephants have reproduced so poorly, in contrast to the population in Addo National Park, just east of the Fynbos Biome. Micronutrient deficiency has been suspected for many years (Carter 1971; Koen et al. 1988). Seydack ( et al . 2000) concluded studies on diet and poor reproductive performance by providing significant evidence that the most probable reason for the rapid decline of the Knysna elephant in recent years despite protection and excellent body condition are attributable to low nutrient/carbon ratios of the diet available to them in the predominantly forest environment to which they were largely confined as a result of residential and agricultural development in the region. (Milewski 2000) suggested that iodine and selenium deficiencies had the greatest effects on reproduction, both elements of which are currently unavailable to the Knysna elephants.

Recently, evidence of photographs of more elephants has emerged (Dep records-2000, 2004; Eggert and Patterson, 2006). However, evidence from monitoring currently reveals the presence of only one animal. Further molecular research is in progress to complement physical monitoring evidence. ORDER: SUIFORMES This order contains one family (Suidae) and species, Potamochoerus larvatus . Studies in the Knysna forest (Seydack, 1990) revealed home ranges of between 4 and 10 km2. Population densities of 0,3 to 0,5 bushpig/km2 were determined for the Knysna forest but appear subject to substantial fluctuations. The mean distance of daily movement was established as 3,0 km (range: 0,48-5,84 km). Activity may occur at any time of the day, with a main peak between 18:00 and 24:00. In the day dense cover is used for resting. Three or four young are usually born after a gestation period of approximately four months, between October and January, in nests, in hollow trees or crevices in wooded krantzes. Seydack (1983) performed an age assessment of the bushpig and found that full-grown animals are two to three years old. ORDER: RODENTIA This is the largest order with four families and eighteen species, one of which is endangered namely, Mystromys albicaudatus (White-tailed rat). As with most of the smaller mammals, very few studies have been undertaken. Family: Muridae Grammomys dolichurus (Woodland thicket rat) This species was recorded in the southern Cape forest by Prof. A.K. Lee (University of Natal:

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Zoology). According to Davis (1974), this record provided an extension to the known distribution of the species.

According to Seydack (1977), small mammal counts in the Goudveld area are low with regard to population densities as well as species diversity. Koen (1991) investigated the effect of rodent granivory on recruitment of the irregularly fruiting Podocarpus falcatus in the Southern Cape. The major granivore was identified as Myomyscus verreauxi (Verreaux's mouse). No significant correlations wee found between the density of seed on the ground and the number of seeds destroyed. Koen (1991) concluded that this species forages randomly and not in a density dependent manner. Additional surveys are urgently needed. ORDER: WHIPPOMORPHA This order consists of six families and fourteen species, three of which are vulnerable and one species near threatened. Thirteen of the species (92.8%) are CITES listed.

Shore-based observations along the coast by Saayman et al. (1972) suggested that humpback dolphins Sousa chinensis formed small schools (c. 7 individuals), fed on nearshore reef fish and invertebrates and remained in the area throughout the year. Bottle-nosed dolphins Tursiops truncates formed intermediate size schools (c. 140 individuals), were abundant during spring and summer, and appeared capable of exploiting both reef and off-shore pelagic food sources. Pelagic dolphins Delphinus delphis and Stenella coeruleoalba formed large schools (c. 670 individuals) and were only seen far offshore. The numbers of southern right whales ( Eubalaena australis ) that frequent the Tsitsikamma coast are low compared to that of the southwestern Cape (Best 2000).

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APPENDIX E

GRNP ANIMAL SPECIES LIST

TERRESTRIAL INVERTEBRATES

Species Life form Area Identified by Stick insect Petrus Brand Per Dr R. Toms [at Transvaal Museum] Acanthodelta distriga Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Acherontia atropos Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Acraea horta Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Acraea sp. Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Acrasia crinita Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Acripia semiviridis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Adicocrita discerpta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Adicocrita koranata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Aeropetes tulbaghia Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Aethaloessa floridalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Aethiopsestis austrina Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Afrodromia sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Afroideus cruciatus Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Afrosteles distans Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Agdistis sp. Moth, Millwood/ Petrus Brand [Identification by Dr D Kroon] delicate Aglossosia flavimarginata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Agrius convolvuli Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Agrotis cinctithorax Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Agrotis segetum Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Alelimma grisea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Aletia viettei Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Identification by the Department of Allodape ceratinoides Harkerville Zoology, University of Witwatersrand Identification by the Department of Allodape exoloma Harkerville Zoology, University of Witwatersrand Identification by the Department of Allodape skaiforem Harkerville Zoology, University of Witwatersrand Identification by the Department of Allodape tridentipes Harkerville Zoology, University of Witwatersrand Identification by the Department of Allodapula melanoplus Harkerville Zoology, University of Witwatersrand Identification by the Department of Allodapula variegata Harkerville Zoology, University of Witwatersrand Aloeides clarki Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81

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Aloeides aranda Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Aloeides quickelbergi Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Amauris albimaculata Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Amauris echeria echeria Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Amazonides ustula Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ambia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ancylolomia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Antanartia hippomene hippomene Millwood Anthophora madecassa Bee Millwood/ Petrus Brand Von Breit. Antiophlebia bracteata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Anua selenaris Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Apis caffra Bee Millwood/ Petrus Brand Von Breit. Apis mellifera vulgaris Bee Millwood/ Petrus Brand Von Breit. Apochrysa leptalea Lacewings Millwood/ Petrus Brand [Identification by Dr M. Mansell, at ARC - Plant Protection Research Institute] Apomyelois bicolorata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Appana sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Argyrographa moderata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Argyrophora arcualis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Argyrophora trophonia Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Asthenotricha inutilis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Athetis sp. Moth Millwood Athetis albipuncta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Athetis cf. Moth Petrus Brand [Identification by Dr D Kroon] Athetis nigra Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Athetis pigra Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Athetis sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Athetis tenebrata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Atractoceros sp. Beetle Millwood/ Petrus Brand [Identification by Dr D Kroon] Automolis benitensis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Batracomomorphus subolivaceous Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Belenois gidica Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Belenois thysa thysa Butterfly Millwood/ Petrus Brand Migdol Belenois zochalia zochalia Butterfly Millwood/ Petrus Brand Migdol Belenois aurota Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Belenois creona severina Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Bicyclus safitza safitza Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Blenina quadripuncta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Boerias maculata Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Bombycomorpha spp Millwood/ Petrus Brand Von Breit. Bombycopsis capicola Eggar Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Borolia ustata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Botys albialis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Brevipecten cornuta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Capys alpheus alpheus Millwood/ Petrus Brand 20/11/5/1 folio 81 Cassionympha cassius Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Catacroptera cloanthe cloanthe Millwood/ Petrus Brand Migdol

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Catochria catocaloides Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Celama iridescens Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Celama megasema Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Celama tineoides Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Chabula quadrinalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Charaxes pelias Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Charaxes karkloof capensis Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Charaxes karkloof trimeni Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Charaxes varances Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Charaxes xiphares xiphares Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Chiasmia streniata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Chlorissa articulicornis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Chloroclystis derasata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Chlorocoma didita Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Chrysoperla pudica Lacewings Millwood/ Petrus Brand [Identification by Dr M. Mansell, at ARC - Plant Protection Research Institute] Chrysoritis cottrelli Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Circulifer sp. Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Cleora betularia Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Cleora tulbaghata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Coelonia mauritii Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Cofana spectra Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Colias electo Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Colotis antevippe gavisa Butterfly Millwood/ Petrus Brand Migdol Colotis evagore antigone Butterfly Millwood/ Petrus Brand Migdol Comostolopsis capensis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Comostolopsis rubristicta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Conservula minor Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Conservula sinisigna Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Copris sp. Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Corgatha producta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Crambus sparsellus Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Craniophora paragrapha Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Craterestra definiens Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ctenoplusia limbirena Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ctenosia psectriphora Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Culladia serranella Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Cyana marshalli Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Cymathoe sp. Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Cymothoe alcimeda Butterfly Millwood/ Petrus Brand Migdol Cymothoe alcimeda alcimeda Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Dagama capensis Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Danaus chrysippus Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Derrioides villaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon]

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Diasemia monostigma Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Dicranuropsis vilis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Dieuches sp. Seed Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Dira clytus clytus Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Dira oxylus Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Dixeia sp. Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Dolichocephala fuscilanx Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Drepanogynis arcuifera Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Drepanogynis mixtaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Drymodromia trochanterata Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Duponchelia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Durbaniella clarki Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Dysaletria sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Earias biplaga Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ectropis spoliataria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Egnasia vicaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eilema caffrana Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eilema sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Empusada chrysota Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Epiplema inconspicua Moth (very Millwood/ Petrus Brand [Identification by Dr D Kroon] small) Erioderus hirtus Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Eublemma sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eublemma sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eublemmistis chlorozonea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euchromius sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eudalaca sp. Moth, m- Millwood/ Petrus Brand Pers Obs. J.Baard large Eumicremma minima Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eupithesia festiva Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eupithesia infelix Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euplexia augens Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euproctis crocata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euproctis haemodetes Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euproctis sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euproctis terminalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eurosia fuliginea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eurosia lineata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Eurytela hiarbas Butterfly Millwood/ Petrus Brand Pers Obs. J.Baard Eustrotia loxosema Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Euzophora villora Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Exilisia bipuncta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Exitianus taeniaticeps Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Eyralpenus testacea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Gaana viridella Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Galleria mellonella Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Gegenes hottentota hottentota Butterfly Millwood

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Gryllotalpa vulgaris Mole cricket Millwood/ Petrus Brand Von Breit. Heliothis [Helicoverpa] Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Hemerobius nairobicus Lacewings Millwood/ Petrus Brand [Identification by Dr M. Mansell, at ARC - Plant Protection Research Institute] Hemerodromia sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Hemerodromia cf. alida Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Heterographis mesosema Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Heteroplema dependens Moth (very Millwood/ Petrus Brand [Identification by Dr D Kroon] small) Heterorachis despoliata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Heterorachis disconotata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Hilara crassitibia Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Hippotion celerio Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Holocerina smilax Emperor Millwood/ Petrus Brand [Identification by Dr D Kroon] moth Horisme minuata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Hypacostemma viridissima Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Hypagoptera rufeola Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Hypena sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Hypoplexia conjuncta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ilemodes heterogyna Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Italochrysa amplipennis Lacewings Millwood/ Petrus Brand [Identification by Dr M. Mansell, at ARC - Plant Protection Research Institute] Junonia archesia Butterfly Millwood/ Petrus Brand Migdol Laelia octophora Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Laelia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lamachella marginalis Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Lamprosiella eborella Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Larentia bitrita Moth Petrus Brand [Identification by Dr D Kroon] Larentioides cacothemon Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Latoia albicosta Slug moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lepidochrysops asters Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Lepidochrysops braueri Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Lepidochrysops robertsoni Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Leto venus Moth, m- Millwood/ Petrus Brand [Identification by Dr D Kroon] large Leucoptera sp. Moth, clothes Millwood/ Petrus Brand [Identification by Dr D Kroon] Leucotrachea cf. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Leucotrachea melanoleuca Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lithacodia normalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Locris sanguinipes Cycada Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Loryma sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Loxostege zaide Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lycophotia perirrorata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lymantria kettlewelli Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Lysceia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon]

management plan GRNP Appendices 60

Macropoliana natalensis Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Macropoliana oheffernani Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Macrotoma natala Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Mauna filia Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Maurilia arcuata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Megopis mutica Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Melanthia ustiplaga Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Metahepialus plurimaculata Moth, m- Millwood/ Petrus Brand [Identification by Dr D Kroon] large Metisella metis Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Moma verbenata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Monoplema fumigera Moth (very Millwood/ Petrus Brand [Identification by Dr D Kroon] small) Mylothris chloris agathina Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Myomantis sp Millwood/ Petrus Brand Per Dr R. Toms [at Transvaal Museum] Napotrephes sp. Cycada Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Nepheronia buquetii buquetii Butterfly Millwood/ Petrus Brand Migdol Nezara prunasis Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Nezara viridula Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Nodaria luctuosa Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Nodaria melaleuca Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Nola fovifera Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Nola mesoscota Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Nopia saxaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Noticula nr Seed Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Nudaurelia spp Moth Millwood/ Petrus Brand Von Breit. Nyodes acatharta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Nyodes lutescens Moth Petrus Brand At Rumohra as a larva Nysius sp. Seed Bug Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Oaracta maculata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Obolcola petronaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ocginus lactipennis Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Ochropleura leucogaster Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ochropleura sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Odontopera erebaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Oedicentra albipennis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Oedilepia capensis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Oligia instructa Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Olybama uliginosalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon]

management plan GRNP Appendices 61

Omizodes ocellata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Omocenoides breyeri Slug moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ophiusa ambigua Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pachypasa albofasciata Eggar moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Palpares speciosus Lacewings Millwood/ Petrus Brand [Identification by Dr M. Mansell, at ARC - Plant Protection Research Institute] Palpita unionalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pantoctenia gemmans Slug moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pareclipsis anophthalma Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pareclipsis phaeopis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Paroruza subductata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Parotis sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Parthenodes angularis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pashiodes mesoleucalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pasteosia irrorata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pelecystola melanchares Moth, clothes Millwood/ Petrus Brand [Identification by Dr D Kroon] Penisa albigrisea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Peridroma sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Identification by the Department of Zoology, University of Peripatopsis clavigera Harkerville Witwatersrand Identification by the Department of Zoology, University of Peripatopsis sedgwicki Harkerville Witwatersrand Perizoma epipercna Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Perizoma sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Phasis thero Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Philobota erastris Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Phymateus morbillosus Locust Millwood/ Petrus Brand Von Breit. Piercia bryophilaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Piercia cidariata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Piercia ciliata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pingasa distensaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Platypalpus cf. aciculatus Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Platypalpus cf. inermifemur Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Platypalpus cf. manikoosi Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Pleuroptya mesoleucalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pleuroptya nr Moth Petrus Brand [Identification by Dr D Kroon] Poecilmitis chrysaor Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Poecilmitis palmus margaretae Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Polia speyeri Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Princeps dardanus cenea Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Princeps demodocus Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Princeps nireus Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon] Princeps nireus lyaeus Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Problepsis latonaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Protosteira spectabilis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Psara vecordalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pseudaglossa pulverea Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pseudomaenas anguinata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon]

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Pseudonympha magus Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Psilocera pulverosa Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Psilocladia obliquata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pusiola asperatella Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pusiola sordida Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Pusiola sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Rhadinomphax nr Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Rhaphium currani Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Rhesala / Rhesalides sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Rhizoplatys auriculatus Beetles Millwood/ Petrus Brand Identification by Dr M Krüger of the Transvaal Museum Rhodogastria amasis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Rhodometra sacraria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Rhypopteryx rhodalipha Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Ribistella cf. submaculata Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Rubistella bimaculata Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Rubistella sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Sarimarais peringueyi Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scalidomia horidella Moth, clothes Millwood/ Petrus Brand [Identification by Dr D Kroon] Schistissa uniformis Monkey moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scoparia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scopula acentra Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scopula inscriptata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scopula instructata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scopula sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scotopteryx albiclausa Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scotopteryx crenulimargo Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scotopteryx ferridotata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Scotopteryx subrectiaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Selidoris deligata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Seratosolen (f) capensis Wasp Millwood/ Petrus Brand Pers Obs. J.Baard Seratosolen (m) capensis Wasp Millwood/ Petrus Brand Pers Obs. J.Baard Sesamia sp. Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Siccia caffra Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Sicyodes costipicta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Somatina centrophora Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Stathmopoda nr Petrus Brand [Identification by Dr D Kroon] Stenoproctus sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Stygionympha vigilans Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Sychophaga cyclostigma Wasp Millwood/ Petrus Brand Pers Obs. J.Baard Sympycnus cf. caffer Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Sympycnus plexsim Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Syngatha purpurascens Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Syntormoneura munroi Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Tachypeza sp. Fly Millwood/ Petrus Brand 20/11/5/1 of 27/05/92 Taviodes subjecta Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Tegulifera obovalis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Temnora orpheus Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Temnora plagiata Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Temnora pylades Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Temnora zantus Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon]

management plan GRNP Appendices 63

Tetracme truncataria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Theretra capensis Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Theretra orpheus Hawkmoth Millwood/ Petrus Brand [Identification by Dr D Kroon] Thestor braunsi Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Thestor murrayi Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Thyretes hippotes Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Tinea spilocoma Moth, clothes Millwood/ Petrus Brand [Identification by Dr D Kroon] Tracheplexia albimacula Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Trichoplusia chalcedona Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Triphassa stalachtis Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Vanessa cardui Butterfly Millwood/ Petrus Brand 20/11/5/1 folio 81 Vietteania torrentium Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Virachola antalus Butterfly Millwood/ Petrus Brand Migdol Xanthorhoe melissaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Xanthorhoe transjugata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Xenimpia maculosata Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Xestocephalus sp. Lacewings Millwood/ Petrus Brand [Identification by Michael Stiller, at ARC - Plant Protection Research Institute] Xylopteryx protearia Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Yponomeuta Moth, Millwood/ Petrus Brand [Identification by Dr D Kroon] colourfull Zamarada transvisaria Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Zitha Moth Millwood/ Petrus Brand [Identification by Dr D Kroon] Zizeeria knysna Butterfly Millwood/ Petrus Brand [Identification by Dr D Kroon]

management plan GRNP Appendices 64

REPTILES

SARDB Order Suborder Family Scientific English Name Status IUCN Status Chelonii Cryptodira Cheloniidae Caretta caretta loggerhead turtle Vulnerable Endangered (A1abd) Chelonii Cryptodira Cheloniidae Chelonia mydas green turtle Vulnerable Endangered (A2bd) Critically Endangered Chelonii Cryptodira Cheloniidae Eretmochelys imbricata hawksbill sea turtle Vulnerable (A1bd) Critically Endangered Chelonii Cryptodira Dermochelyidae Dermochelys coriacea leatherback sea turtle Vulnerable (A1abd) Chelonii Cryptodira Pelomedusidae Pelomedusa subrufa marsh terrapin Null Null Chelonii Cryptodira Testudinidae Chersina angulata angulate tortoise Null Null Chelonii Cryptodira Testudinidae Geochelone pardalis leopard tortoise Null Null Chelonii Cryptodira Testudinidae Homopus areolatus parrot-beaked tortoise Null Null Squamata Sauria Agamidae Agama atra atra southern rock agama Null Null Squamata Sauria Chamaeleonidae Bradypodion damaranum Knysna dwarf chameleon Null Null Squamata Sauria Cordylidae Chamaesaura anguina anguina Cape grass lizard Null Null Squamata Sauria Cordylidae Cordylus coeruleopunctatus blue-spotted girdled lizard Null Null Squamata Sauria Cordylidae Cordylus cordylus Cape girdled lizard Null Null Pseudocordylus microlepidotus Squamata Sauria Cordylidae microlepidotus Cape crag lizard Null Null Squamata Sauria Gekkonidae Afrogecko porphyreus marbled leaf-toed gecko Null Null Squamata Sauria Gekkonidae Pachydactylus bibronii Bibron's Gecko Null Null Squamata Sauria Gekkonidae Pachydactylus geitje Ocellated gecko Null Null Squamata Sauria Gekkonidae Pachydactylus maculatus spotted gecko Null Null Squamata Sauria Gerrhosauridae Tetradactylus seps short-legged seps Null Null Squamata Sauria Gerrhosauridae Tetradactylus tetradactylus common long-tailed seps Null Null Squamata Sauria Gerrhosauridae Gerrhosaurus flavigularis yellow-throated plated lizard Null Null Squamata Sauria Gerrhosauridae Tetradactylus africanus African Long-tailed Seps Null Null Squamata Sauria Lacertidae Nucras lalandii Delalande’s sandveld lizard Null Null Pedioplanis lineoocellata Squamata Sauria Lacertidae pulchella spotted sand lizard Null Null Squamata Sauria Lacertidae Tropidosaura gularis Cape mountain lizard Null Null Tropidosaura montana Squamata Sauria Lacertidae montana common mountain lizard Null Null Squamata Sauria Scincidae Acontias meleagris meleagris Cape legless skink Null Null Squamata Sauria Scincidae Eremias lineocelata Sand lizard Null Null Squamata Sauria Scincidae Mabuya capensis Cape Skink Null Null Squamata Sauria Scincidae Mabuya homalocephala smithii Smith’s skink Null Null Varanus exanthematicus Squamata Sauria Varanidae albigularis Rock Monitor Null Null Squamata Serpentes Colubridae Amplorhinus multimaculatus many-spotted snake Null Null Squamata Serpentes Colubridae Crotaphopeltis hotamboeia herald snake Null Null Squamata Serpentes Colubridae Dasypeltis scabra common egg-eater Null Null Squamata Serpentes Colubridae Dispholidus typus typus boomslang Null Null Squamata Serpentes Colubridae Duberria lutrix lutrix common slugeater Null Null Squamata Serpentes Colubridae Lamprophis aurora Aurora house snake Null Null Squamata Serpentes Colubridae Lamprophis capensis brown house snake Null Null Squamata Serpentes Colubridae Lamprophis inornatus olive house snake Null Null

management plan GRNP Appendices 1

Squamata Serpentes Colubridae Lycodonomorphus rufulus common brown water snake Null Null Squamata Serpentes Colubridae Lycophidion capense capense Cape wolf snake Null Null Squamata Serpentes Colubridae Philothamnus hoplogaster green water snake Null Null Philothamnus natalensis Squamata Serpentes Colubridae occidentalis eastern green snake Null Null Prosymna sundevallii Squamata Serpentes Colubridae sundevallii Sundevall’s shovel-snout Null Null Squamata Serpentes Colubridae Psammophis crucifer cross-marked grass snake Null Null Squamata Serpentes Colubridae Psammophylax rhombeatus spotted skaapsteker Null Null Squamata Serpentes Colubridae Pseudaspis cana mole snake Null Null Squamata Serpentes Elapidae Hemachatus haemachatus Rinkhals Null Null Squamata Serpentes Elapidae Homoroselaps lacteus spotted harlequin snake Null Null Squamata Serpentes Elapidae Naja nivea Cape cobra Null Null Squamata Serpentes Elapidae Pelamis platurus yellow-bellied sea snake Null Null Squamata Serpentes Leptotyphlopidae Leptotyphlops nigricans black thread snake Null Null Squamata Serpentes Typhlopidae Rhinotyphlops lalandei Delalande ’s Beaked Blind Snake Null Null Squamata Serpentes Viperidae Bitis arietans arietans puff adder Null Null Squamata Serpentes Viperidae Bitis atropos berg adder Null Null Squamata Serpentes Viperidae Causus rhombeatus common night adder Null Null

FROGS

CITES NEMBA Priority Order Family Scientific English Name Redlist 2003 IUCN 2000 Listing Category Species Anura Bufonidae Vandijkophrynus Cape sand toad Least concern NL NULL NULL

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angusticeps Vandijkophrynus Anura Bufonidae gariepensis Karoo toad Least concern NL NULL NULL Anura Bufonidae Amietophrynus pardalis Eastern leopard toad Least concern NL NULL NULL Anura Bufonidae Amietophrynus rangeri Raucous toad Least concern NL NULL NULL Tradouw mountain Anura Bufonidae Capensibufo tradouwi toad Least concern NL NULL NULL X Anura Heleophrynidae Heleophryne regis Southern ghost frog Least concern NL NULL NULL X Knysna leaf-folding Anura Hyperoliidae Afrixalus knysnae frog Endangered Endangered NULL Endangered X Anura Hyperoliidae Hyperolius horstockii Arum lily frog Least concern NL NULL NULL X Anura Hyperoliidae Hyperolius marmoratus Painted reed frog Least concern NL NULL NULL X Anura Hyperoliidae Semnodactylus wealii Rattling frog Least concern NL NULL NULL Anura Brevicipitidae Breviceps fuscus Plain rain frog Least concern NL NULL NULL X Cape mountain rain Anura Brevicipitidae Breviceps montanus frog Least concern NL NULL NULL X Anura Brevicipitidae Breviceps rosei Rose's rain frog Least concern NL NULL NULL X Anura Pyxicephalidae Cacosternum boettgeri Boettger's caco Least concern NL NULL NULL Anura Pyxicephalidae Cacosternum nanum Bronze caco Least concern NL NULL NULL Anura Pipidae Xenopus laevis Common platanna Least concern NL NULL NULL Anura Pyxicephalidae Amietia angolensis Common river frog Least concern NL NULL NULL Anura Pyxicephalidae Amietia fuscigula Cape river frog Least concern NL NULL NULL Anura Pyxicephalidae Strongylopus bonaespei Banded stream frog Least concern NL NULL NULL X Anura Pyxicephalidae Strongylopus fasciatus Striped stream frog Least concern NL NULL NULL Anura Pyxicephalidae Strongylopus grayii Clicking stream frog Least concern NL NULL NULL Anura Pyxicephalidae Tomopterna delalandii Cape sand frog Least concern NL NULL NULL X

MAMMALS

Order Family Scientific name Common name Endemism Endemism CITES 1986 Rodentia Muridae Acomys subspinosus Cape spiny mouse W Cape WCP END NL

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Afrosoricida Chrysochloridae Amblysomus corriae Fynbos Golden Mole w/e Cape CFK END NL Afrosoricida Chrysochloridae Amblysomus hottentotus Hottentot golden mole SA NL Carnivora Mustelidae Aonyx capensis African clawless otter Appendix II Carnivora Otariidae Arctocephalus pusillus South African Fur Seal Appendix II Carnivora Otariidae Arctocephalus tropicalis Subantarctica Fur Seal Appendix II Carnivora Herpestidae Atilax paludinosus Marsh mongoose NL Whippomorpha Balaenidae Balaenoptera edeni Bryde's whale Appendix I Rodentia Bathyergidae Bathyergus suillus Cape dune molerat W Cape WCP END NL Carnivora Felidae Caracal caracal Caracal Appendix II Primates Cercopithecidae Cercopithecus pygerythrus Vervet Monkey Appendix II Afrosoricida Chrysochloridae Chlorotalpa duthieae Duthie's golden mole w/e Cape NL Eulipotyphla Soricidae Crocidura flavescens Greater red musk shrew NL Rodentia Bathyergidae Cryptomus hottentotus African Mole-rat NL Whippomorpha Delphinidae Delphinus delphis Short-beaked common Dolphin Appendix II Rodentia Muridae Dendromus melanotis Grey Climbing Mouse Rodentia Muridae Dendromus mesomelas Brants' climbing mouse NL Rodentia Muridae Dendromus mystacalis Chestnut climbing mouse NL Macroscelidae Macroscelididae Elephantulus edwardii Cape Rock Elephant-shrew SA SA NL Chiroptera Pteropodidae Epomophorus wahlbergi Wahlberg’s epauletted fruit bat NL Chiroptera Vespertilionidae Eptesicus hottentotus Long-tailed serotine bat NL Whippomorpha Balaenidae Eubalaena australis Southern right whale Appendix I Carnivora Felidae Felis silvestris African wild cat Appendix II Carnivora Viverridae Galerella pulverulenta Cape grey mongoose swestern NL

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Africa Carnivora Viverridae Genetta genetta Small-spotted genet NL South African large-spotted Carnivora Viverridae Genetta tigrina genet NL Rodentia Bathyergidae Georychus capensis Cape Mole-Rat SA NL Rodentia Muridae Grammomys dolichurus Woodland thicket rat NL Whippomorpha Delphinidae Grampus griseus Risso's dolphin NL Rodentia Myoxidae Graphiurus murinus Woodland dormouse NL Rodentia Myoxidae Graphiurus ocularis Spectacled dormouse w/e/n Cape NL Carnivora Herpestidae Herpestes ichneumon Large grey mongoose NL Rodentia Hystricidae Hystrix africaeaustralis Cape porcupine NL Carnivora Mustelidae Ictonyx striatus Striped Polecat NL Chiroptera Vespertilionidae Kerivoula lanosa Lesser woolly bat NL Whippomorpha Kogiidae Kogia breviceps Pygmy sperm whale Appendix II Whippomorpha Kogiidae Kogia sima Dwarf sperm whale Appendix II Lagomorpha Leporidae Lepus capensis Cape hare NL Lagomorpha Leporidae Lepus saxatalis Scrub hare NL Carnivora Phocidae Lobodon carcinophaga Crab-eating seal Proboscidea Elephantidae Loxodonta africana African Elephant Appendix II Macroscelides swestern Macroscelidae Macroscelididae proboscideus Round-eared Elephant-shrew Africa NL Rodentia Muridae Mastomys natalensis Natal Multimammate mouse NL Whippomorpha Balaenopteridae Megaptera novaeangliae Humpback whale Appendix I Carnivora Mustelidae Mellivora capensis Honey Badger Appendix III

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Whippomorpha Ziphiidae Mesoplodon densirostris Blainville's beaked whale Appendix II Whippomorpha Ziphiidae Mesoplodon hectori Hector's beaked whale Appendix II Chiroptera Vespertilionidae Miniopterus fraterculus Lesser long-fingered bat NL Chiroptera Vespertilionidae Miniopterus schreibersii Schreibers' long-fingered bat NL Carnivora Phocidae Mirounga leonina Southern elephant seal Appendix II Rodentia Muridae Mus minutoides Pygmy mouse NL Rodentia Muridae Myomyscus verreauxi Verreaux's mouse w/e Cape NL Eulipotyphla Soricidae Myosorex cafer Dark-footed forest shrew NL Eulipotyphla Soricidae Myosorex longicaudatus Long-tailed forest shrew w/e Cape NE NL Eulipotyphla Soricidae Myosorex varius Forest shrew NL Chiroptera Vespertilionidae Myotis tricolor Temminck's hairy bat NL Rodentia Muridae Mystromys albicaudatus White-tailed rat NL Chiroptera Vespertilionidae Neoromicia capensis Cape serotine bat NL Chiroptera Nycteridae Nycteris thebaica Egyptian slit-faced bat NL Whippomorpha Delphinidae Orcinus orca Killer whale Appendix II Ruminantia Bovidae Oreotragus oreotragus Klipspringer NL Tubulidentata Orycteropodidae Orycteropus afer Aardvark NL Rodentia Muridae Otomys irroratus Vlei rat NL Carnivora Felidae Panthera pardus Leopard Appendix I Primates Cercopithecidae Papio hamadryas Chacma baboon Appendix II Ruminantia Bovidae Pelea capreolus Grey rhebok NL Ruminantia Bovidae Philantomba monticola Blue Duiker Appendix II Whippomorpha Physeteridae Physeter catodon Sperm whale Appendix I

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Chiroptera Vespertilionidae Pipistrellus hesperidus African pipistrelle NL Carnivora Mustelidae Poecilogale albinucha African striped weasel NL Suiformes Suidae Potamochoerus larvatus Bushpig NL Hyracoidea Procaviidae Procavia capensis Rock dassie NL Ruminantia Bovidae Raphicerus campestris Steenbok NL NE WCP and Ruminantia Bovidae Raphicerus melanotis Cape grysbok w/e Cape CFKNE NL Rodentia Muridae Rhabdomys pumilio Four-striped grass mouse NL Chiroptera Rhinolophidae Rhinolophus capensis Cape horseshoe bat w/e/n Cape NL Chiroptera Rhinolophidae Rhinolophus clivosus Geoffroy's horseshoe bat NL Chiroptera Pteropodidae Rousettus aegyptiacus Egyptian rousette NL Rodentia Muridae Saccostomus campestris Pouched Mouse NL Whippomorpha Delphinidae Sousa chinensis Humpback dolphin Appendix II Whippomorpha Delphinidae Stenella coeruleoalba Striped dolphin Appendix II Eulipotyphla Soricidae Suncus infinitesimus Least dwarf shrew NL Ruminantia Bovidae Sylvicapra grimmia Common Duiker NL Chiroptera Molossidae Tadarida aegyptiaca Egyptian Free-tailed Bat NL Chiroptera Emballonuridae Taphozous mauritianus Mauritian Tomb bat NL Ruminantia Bovidae Tragelaphus scriptus Bushbuck NL Whippomorpha Delphinidae Tursiops truncates Bottlenose dolphin Appendix II

management plan GRNP Appendices 7

APPENDIX F

FRESHWATER & ESTUARINE ISSUES AND PROCESSES

HYDROLOGY

Touw system

The catchment area of the Touw system comprises three rivers – Touw River (96.2 km 2), Duiwe River (42.1km 2) and the Langvlei Spruit (8.2km 2) (Hughes & Filmalter 1993, Fijen & Kapp 1995). Virgin MAR into the Touw system has been estimated by Fijen (1995a) as 24.6 x 10 6 m -3. Modelling of flood events emphasised the short residency time of flood waters and hence potential for rapid increases in water level in the lower lakes, particularly the Touw Estuary (Görgens 1979). Expected water levels (meters amsl) in the Touw Estuary that would result during flood events of different return frequency are 5 yr. = 3.22; 10 yr. = 3.61; 20 yr. = 3.88; 50 yr. = 4.01; 100 yr. = 4.93 (Görgens 1979). CSIR (1981) and CSIR (1982) concluded that maintaining the height of the sand sill at the estuary mouth at between 2.1 m and 2.4 m amsl should prevent flooding of low-lying properties by floods with a return frequency of 1 in 50 years or lower. It was also concluded that although the dredging of the connecting channels would improve the circulation of water throughout the system, it would not significantly reduce the peak water levels.

Boshoff & Palmer (1989) have suggested that the constriction at the Serpentine sluice inhibits flow to the upper lakes in times of flooding, whereas Hall (1985b) has argued that the effect is minimal. The sluice has not been operational since the 1980’s.

Swartvlei system

Rivers draining into the Swartvlei System are the Diep River (98.3 km 2), Klein Wolwe River (17.2 km 2), Höekraal River (111.0 km 2), and Karatara River (101.6 km 2) (Whitfield et al. 1983, Hughes & Filmalter 1993). Estimates of freshwater input into the Swartvlei system are given in CSIR (1978), Whitfield et al. (1983) and Fijen (1995b), with the latter also providing data on current and predicted future abstraction by various user sectors. Total evaporation from the Swartvlei system was estimated by Whitfield et al. (1983) as 12.1 x 10 6 m -3 (cf. 1127 mm y -1) or approximately 20% of the average total annual freshwater input of 66 x 10 6 m -3 (CSIR 1978) indicating that under natural conditions Swartvlei Lake is unlikely to become hypersaline.

Severe floods in the Swartvlei system are infrequent. However, when river floods coincide with periods when the estuary mouth is closed, or even with very high spring tides when the estuary mouth is open, then flooding of adjacent land can be expected, especially in the Sedgefield Extension 1 area (Howard-Williams & Allanson 1979). It has been recommended that for hydrological, ecological and social reasons Swartvlei Estuary mouth be artificially breached at 2.0 m amsl (CSIR 1978, Howard-Williams & Allanson 1979, Whitfield et al. 1983).

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There has been a long standing debate about the impact that the rail-bridge dividing Swartvlei Lake and Swartvlei Estuary has on the movement of water between these two systems. Whitfield et al. (1983) indicated that the rail-bridge does interfere with the flow of water, though added that the influence is confined to the immediate vicinity of the bridge as the main restriction to outflow in the system is the narrow estuary mouth. Mathematical modelling of the water flow regime in the Swartvlei system (CSIR 1978) revealed that the complete removal of the rail-bridge would result in a 40% increase in tidal exchange between the lake and the estuary. This increase in tidal exchange, however, would result in a change in the lake level of only a few millimetres, and would have no discernible effect on the estuary mouth (CSIR 1978). Later modelling exercises provided revised information on flow rates and volumes (Huizinga 1987), though conclusions regarding the low impact of the rail-bridge remained.

Knysna system

The Knysna River runs mainly north south and cut deep narrow incisions into the underlying Table Mountain Sandstone. The Knysna River feeds from two catchments, K50a and K50b. The northern part of K50a hosts Oubos River (4.7km) and two other unnamed rivers. These originate in mountain fynbos largely planted with plantation. The Bobbejaan River (7km) is the first major west-east tributary inside the park. The Kruis River (18.7km) enters below the Bobbejaan River and drains mainly from plantation covered vegetation. Many smaller unnamed tributaries join the Knysna River from both east and west, often from forest, but more often from plantation areas. The next major tributaries are the Lawnwood River (6km) from the west and the Rooiels River (13.9 km) from the east. From the confluence of the Knysna- and Rooiels Rivers, the Knysna River runs between private land on the west and forestry/farmland on the east. The tributaries from the west drains mainly from indigenous forest and to a lesser extent cultivated land.

The Gouna River (21.8km) is the largest tributary of the Knysna River. The Steenbras River (9.4km) is the largest tributary of the Gouna River. Both these rivers drain mainly from indigenous forest and to a much lesser extent, fynbos. The Gouna River and the Steenbras River are both boundaries of SANParks’ land and the most northern rivers in catchment K50b.

Tributaries entering the Knysna River below the confluence of the Knysna and the Gouna Rivers drain from indigenous forest and to a lesser extent farmland to the west of the Knysna River, and plantation to the east of the Knysna River.

A few rivers join the Knysna River in the Knysna Estuary. These rivers run through populated areas, industrial areas and informal settlements and are often of poor quality.

The Knysna River is approximately 64 km long (Grindley 1985). Some confusion appears to exist regarding the size of the Knysna drainage basin which is given as 400 km 2 (Day et al . 1952), 526 km 2 (Noble & Hemens 1978, Day 1981) and 525 km 2 (Pitman et al . 1981), of which the Knysna River basin comprises 426 km 2 (Noble & Hemens 1978), 337 km 2 (Zwamborn 1980) or 315 km 2 (Reddering & Esterhuysen 1984). Presumably the latter two areas exclude the estuary and local drainage (7 streams) which cover approximately 100 km 2 (Noble & Hemens 1978, Day 1981). Reddering (1994) descries each of 12 sub-catchments draining into the estuary in terms of size, vegetation cover, soils, development and sediment yield. The Knysna estuary is an S-shaped stretch of water, 1633 ha in extent (Duvenhage

management plan GRNP Appendices 9

1983); with a channel approximately 19 km long and up to 2 km wide. It has a tidal reach of about 17 km (Reddering & Esterhuysen 1984).

Flow in the Knysna River is perennial (Reddering & Esterhuysen 1984), with virgin MAR estimated by Noble & Hemens (1978) as 110 x 10 6 m 3 y-1 , and by Pitman et al . (1981) as 133 x 10 6 m 3 y-1. Haw (1984) however, gives MAR at Charlesford Farm as only 61 x 10 6 m 3 y-1. Day et al . (1952) maintain that floods occur, on average, once every 10 to 12 years, at which time the estuary is heavily stained by inflowing sediments. Modelling of flows in Knysna Estuary have been undertaken to assess the effect of flood water on water height (CSIR 1976) and in some cases on proposed developments (CSIR 1974). Run-off frequency curves for catchments K5M01and K5M02 are presented by Hughes & Görgens (1981). The Gouna River (K5M01) generally has lower coefficients of variation that the other as this catchment has lower slopes and less relief than mountain catchments.

The highest flood-level recorded in the estuary between 1965 and 1985 was MSL + 2.0m (Grindley 1985). MSL is given by Grindley as 1.16m, which would equate to a water level in the estuary of +3.16m. Marker (2000) maintains that during a storm event on 16 June 1996 that was accompanied by extreme low pressure at spring tide, water levels at Thesen Jetty reached the highest level on record, 0.20m above maximum spring-tide. Furthermore surges at 15-minute intervals added between 0.10 and 0.05 m to the maximum water level. Although Marker (2000) does not give the actual recorded water height, using values of HAT reported by Grindley (1985) (2.29m), water levels in the estuary would have been between 2.54 and 2.59m amsl.

Largier et al. (2000) described three hydrographic regimes in the estuary created by the movement of river and ocean waters, and descried the effects of water through-flow, circulation and diffusion on spatial and temporal variability in salinity and water temperature. The bay regime (extending inland from the estuary mouth between 3km during neap tides and 7.5km during spring tides) is well flushed by tidal flows and exhibits salinities and temperatures similar to the ocean. The lagoon regime (extending inland from the upper reaches of the bay regime and seldom beyond White Bridge) is flushed less rapidly, subject only to tidal diffusion effects, and exhibits long residence times. Salinities here are close to ocean salinity, but temperatures may be several degrees warmer. The estuary regime (extending inland from the upper reaches of the lagoon regime) is characterised by the effects of river inflow and stratification, and is fairly well flushed by density-driven estuarine circulation. The lagoon regime expands as the estuary regime shrinks in response to decrease in river flow and as the bay regime shrinks in response to decrease in tidal range (Largier et al. 2000). The need for sustained river flow to maintain estuarine conditions in the sector above the White Bridge and to allow some variation in salinity in the lagoon regime has been stressed. If the present retention time of some 20 days in the estuarine regime is increased to greater than 50 days as a result of decreased freshwater inputs, water quality will deteriorate (Largier et al. 2000).

Haw (1984) assessed the effect of reduced freshwater inflows into Knysna Estuary on salinity, with cessation of flow resulting in salinities up to 36 g kg -1 in the upper reaches. Modelling of the effects of freshwater flow reductions and flood on salinities was undertaken by Huizinga (1985). Reduced freshwater results in an increase in salinity in the upper reaches of the estuary, with the effects of minor floods (peak 40 m 3 s -1) on salinity reduction visible for about 10 days, and the effects of major floods (peak 328 m 3 s -1) visible for about 15 days, with at maximum flood the salinity throughout the estuary dropping to almost 0 mg kg -1 (Huizinga 1985).

management plan GRNP Appendices 10

Schumann (2000) used temperature variability to demonstrate the hydrodynamics of the estuary, particularly in terms of ocean-estuary exchanges. Ocean temperatures were generally found to be colder than estuary temperatures, with variations in water temperature as a result of tidal fluctuations clearly evident. The influence of cold-water upwelling on estuary temperature was demonstrated in February 1998 where a temperature drop of over 13 °C occurred in about 2 hours.

Tsitsikamma systems

Thirteen perennial rivers flow southward through the Tsitsikamma region of the park (see map below showing hydrology). The largest of these rivers are the Groot River (West), the Bloukrans-Vark, Storms-Witteklip and Elands-Kruis systems, which have calculated catchment areas of 87, 80, 98 and 82 km² respectively (Morant and Bickerton 1983; Chief Director of Surveys and Mapping, 1979, 1980, 1981). The delegation of DWAF managed State Forest land in the southern Cape to SANParks in 2005, allows SANParks greater jurisdiction over the catchment of tributaries of the Elands-, Storms-, Elandsbos-, Bloukrans- and Groot (West) rivers. Soetkraal ‘encompasses’ the upper catchment of Langbos, Palmiet and Diep rivers (Russell 2002).

Figure 4. Hydrology of the GRNP

PHYSICAL AND CHEMICAL PROCESSES

Touw system

Temperature variations in the lakes and estuaries of the Touw system follow a seasonal pattern, generally ranging between 10-14°C in winter and 25-29°C in summer (Whitfield et al . 1983; Russell 1996b). The salinity of the lakes increases the further they are removed from the sea, with Eilandvlei averaging between 6 and 10 g kg -1, Langvlei 10 and 13 g kg -1, and Rondevlei between 12 and 16 g kg -1 (Whitfield et al. 1983; Russell 1999a). An analysis of the salt budget of the Touw system suggests that the Serpentine channel dampens tidal processes management plan GRNP Appendices 11 to the upper lakes to such an extent that they play only a minor role in the overall budget (Hall 1985b). Furthermore, the narrow channel between Eilandvlei and Langvlei prevents any tidal salt input to the upper lakes (Hall 1985b). Turbidities recorded in the Wilderness lakes and estuaries are moderate, with the average of all waterbodies being below 10 NTU (Whitfield et al 1983; Russell 1999a). The only recorded incidence of low oxygen concentrations resulting in the death of organisms occurred in Rondevlei during March 1993 when the senescence of a dinoflagellate/algal bloom resulted the mean oxygen concentration in Rondevlei declining to below 1 ppm, which resulted in the death of several large Cape stumpnose ( Rhabdosargus holubi ) and white steenbras ( Lithognathus lithognathus ) (Russell 1994).

The importance of the sea as a source of P in the Touw River Estuary has been emphasised by Allanson & Whitfield (1983), who, from a study undertaken between January 1979 and June 1981, report an increase in TP from 9 to 17 g l -1 following the breaching of the bar. SRP concentrations however remained low, not exceeding 2.2 g l -1. Similar concentration of P- ions have been recorded in Eilandvlei during non-flood condition, whereas in Langvlei and Rondevlei, TP levels as high as 66 and 117 g l -1 respectively have been recorded (Allanson & Whitfield 1983). Despite these comparatively high levels of TP, only occasionally have SRP levels up to 20 g l -1 been recorded in Langvlei and Rondevlei, which is the point at which, if all other factors are propitious, marked algal growth could occur (Allanson & Whitfield 1983). NO 3-N concentrations in the Touw River Estuary between 1979 and 1981 were found to vary between 7 and 60 g l -1 (Allanson & Whitfield 1983) which is well below that which could be considered to constitute serious organic pollution. Similar low levels of NO 3-N were recorded in the other waterbodies of the Touw system during non-flood periods, with the exception of Langvlei, where values as high as 569 g l -1 have been recorded. However, as Allanson & Whitfield (1983) point out, elevated levels of plant nutrients in the Wilderness lakes were found to occur contemporaneously with changes in the use of the floodplain by large numbers of waterbirds. Surveillance of water chemistry in the Touw system during flood periods has demonstrated that the inflowing streams are an important source of both N- and P-ions into the system (Allanson & Whitfield 1983). Elevated concentrations of both NO 3-N and PO 4-P have been recorded in the Touw River Estuary, Eilandvlei and Langvlei during flood periods, whereas the concentration of N- and P-ions in Rondevlei, with no feeder streams, remained relatively unaltered (Allanson & Whitfield 1983).

A sediment sample collected adjacent to the yacht club in Eilandvlei during April 1977 (Watling 1977) was found to have levels of manganese above those expected when compared with samples collected in Rondevlei and Langvlei. A core sample taken in this locality also showed an overall elevation in values for copper, lead, zinc, cobalt, nickel, cadmium and chromium, with it being hypothesised that these levels represent some degree of anthropogenic contamination (Watling 1977).

Investigations of the concentration of chlorinated hydrocarbon, PCB and Dieldrin residues in the body tissues of aquatic birds in the Touw system have yielded conflicting results. In 1983 De Kock & Boshoff (1987) detected low concentrations of t-DDT and PCB in the body tissues of reed cormorant, darter, whitebreasted cormorant, blacknecked grebe, great crested grebe and grass owl. Similarly, a fish eagle egg, collected at Sedgefield during 1985 had a low concentration of t-DDT (0.21 g g -1 wet weight), and no PCBs or Dieldrin residues (De Kock

management plan GRNP Appendices 12

& Lord 1986). In contrast, in 1984 and 1985 De Kock & Simmons (1988) recorded high levels of Dieldrin (1.89 g g -1) and t-DDT (6.26 & 5.08 g g -1) in African marsh harrier eggs from the Touw system, with low ratios of DDE/t-DDT in several eggs indicating recent inputs of DDT into the environment.

Swartvlei system

Temperature variations in the Swartvlei systems follow a seasonal pattern, generally ranging between 10-14°C in winter and 25-29°C in summer (Whitfield et al . 1983; Russell 1996b). Whitfield et al . (1983) maintain that closure of the Swartvlei Estuary mouth has little effect on water temperature, though during summer when the estuary mouth is open, the waters near the mouth are generally 1°C cooler than further up the estuary. Water temperatures as high as 32°C have been recorded in Swartvlei Estuary (Russell 1996b).

Swartvlei Lake is normally meromictic, as a result of its usual stratification into water layers of different densities, caused by vertical stratification in salt concentration (Robarts & Allanson 1977; Howard-Williams & Allanson 1978). The bottom layer of high salinity water can be up to 5 m thick (Whitfield et al . 1983). Stratification prevents wind from mixing the oxygenated surface water with those below, thus decomposition of organic matter in the bottom sediments rapidly uses up the oxygen and increases the concentration of carbon dioxide (Howard-Williams & Allanson 1979) making the bottom half of Swartvlei Lake a very toxic environment for animal life. When Swartvlei Estuary mouth is closed, wind mixing of the surface waters of the lakes gradually breaks down the salinity layering (Robarts & Allanson 1977; Allanson & Howard-Williams 1984). The longer the mouth remains closed the more likely it is that stratification will break down, with resultant oxygenation of the lower waters of the lake.

Water clarity undergoes periodic dramatic reductions which is generally associated with flood conditions (Whitfield et al . 1983; Russell 1999a). Barker (1985) measured TSS concentrations as high as 4 590.0 mg l -1 in the Klein Wolwe River during a flood event in June 1984. The consequence of these periodic massive inputs of sediment into the lakes was recorded in Swartvlei Lake in the early 1980s where there was an approximate 60% decline in primary production (Whitfield 1982), largely due to a reduction in the standing crop of submerged macrophytes as a consequence of a reduction in light penetration (Whitfield 1982; Allanson & Howard-Williams 1984).

Horizontal variations in dissolved oxygen in the Swartvlei system depend largely on the distribution of aquatic plants, with higher oxygen values being associated with the presence of aquatic plants (Howard-Williams & Allanson 1979). The closing of the estuary mouth has no effect on the mean dissolved oxygen values in Swartvlei Estuary (Howard-Williams & Allanson 1979). Deoxygenation has been recorded in localised areas in the Swartvlei Estuary towards the end of the open phase (Howard-Williams & Allanson 1979). These areas are at the sides of the channel where mats of floating algae start to rot, and in deeper portions of the estuary.

management plan GRNP Appendices 13

The production of hydrogen sulphide (H 2S) gas by anaerobic decomposition of organic matter and the resultant ‘rotten egg’ odour has been discussed by Russell (1997).

Soluble reactive phosphate (SRP) (PO 4-P) concentration in Swartvlei Lake is very low (± 1 g l-1) and is often present in undetectable quantities (Whitfield et al . 1983). It is thought that dissolved humic matter from the rivers plays a significant role in binding phosphate, and hence contributing to the nutrient poor status of the waterbody (Howard-Williams 1977). Total dissolved phosphorous (TDP) values range from 10 to 20 g l -1, and total phosphorous (TP) up to 30 g l -1 (Whitfield et al . 1983). SRP, TDP and TP in the stratified bottom waters of the lake all exceed 100 g l -1 (Howard-Williams 1977) but are unavailable for plant growth (Howard-Williams 1977). The oligotrophic status of Swartvlei Lake has a number of biological consequences, which include low phytoplankton primary production (Robarts 1976), low zooplankton biomass (Coetzee 1981a) and low ichthyoplankton densities (Whitfield 1989b). TDP concentrations in Swartvlei Estuary remains fairly constant at about 24 g l -1 (Whitfield et al . 1983), though increases in concentration (up to 260 g l -1) have been recorded in deoxygenated saline areas. The main source of P for the estuary is the sea, and during the tidal phase there being a net import and accumulation of P in the estuary (Liptrot 1978; Whitfield et al . 1983). An extensive net outflow of P occurs from Swartvlei Estuary during the strong outflow phases immediately after the estuary mouth is opened. Most of the P leaving the estuary is in particulate form, evidenced by the abundance of estuarine plants which usually litter surrounding beaches shortly after the estuary is opened. The concentration of N-ions in Swartvlei Estuary are generally low, with concentrations in 1976 being found to -1 range from not detectable to 21 g l [NO 2+NO 3]-N (Coetzee 1978). Howard-Williams and -1 Allanson (1979) recorded values for nitrate (NO 3-N) ranging from 3 to 30 g l , and Robarts -1 (1973) recorded values for ammonia (NH 4-N) ranging from not detectable to 4 g l .

The distribution of trace elements (copper, lead, zinc, iron, manganese, cobalt, nickel, chromium, cadmium, mercury, magnesium, aluminium, sodium, calcium, strontium, potassium) in water samples, surface sediments and cores was investigated by Watling (1977). In this study it was found that the concentration of trace metals in water samples were generally low, with the exception of iron and manganese in Swartvlei Lake. High levels of nickel from sediments in Swartvlei Estuary may represent contamination from adjacent residential areas. Watling (1977) also suggests that a significant increase in the concentration of lead (and in several instances the concentration of other trace metals) in the upper portions of cores from the Swartvlei system indicate possible pollution from motor fuels or paint.

Knysna system

Ranges of the water quality parameters water temperature, salinity, dissolved oxygen, pH and secchi disk depth in Knysna Estuary from 1990 to 1994 are given in Russell (1996b) and compared with earlier data as given by Day et al . (1952), Day (1967, 1981), Grindley & Eagle (1978) and Haw (1984). It was concluded that no clear long-term changes in recorded water quality parameters were evident. Allanson et al . (2000) undertook comparisons of secchi disk depth with earlier studies in 1991-1994 (Russell 1996b) and 1947 (Day 1967), and concluded that no significant change had occurred in water clarity. Nevertheless the potential of elevated

management plan GRNP Appendices 14 sediment inflow into the estuary as a result of runoff from quarries and disturbed catchment areas (Reddering 1994) remain an environmental concern (Russell 1996b, Allanson et al . 2000), particularly with regard to overall water transparency, the growth of intertidal sandbanks, and the disruption of nutrient pathways for filter feeders.

Allanson et al . (2000) provide values for water quality indicators (pH, TSS, chlorophyll-a, ammonium, nitrate+nitrite, soluble reactive phosphate), measured in Knysna Estuary during 1996 and 1997. In terms of the input of plant nutrients it was concluded that dilution and dispersion by tidal flow is sufficient to prevent overt signs of enrichment either through excessive phytoplankton growth or accumulation of floating macroalgae (Allanson et al . 2000), contrary to the earlier findings of Grindley & Snow (1983). In general the concentration of chlorophyll-a is low, and the water column is oligotrophic. Changes in nutrient levels are closely allied to specific events such as river flooding and sewage treatment plant effluent and/or stormwater inflows from surrounding urban areas. Allanson et al (2000) found there to be a qualitative improvement in chemical water quality when compared to data reported by Grindley & Eagle (1978), thought to be due to an improvement in flow around Thesens Island and an upgrading of the sewage purification works.

The potential for pollution of the Knysna Estuary is high, with in excess of 111 drainage pipes, drainage ditches and culverts discharging storm-water from residential and industrial areas directly into the estuary. In the Ashmead channel, in the region of the overflow from the Knysna sewage treatment works, elevated nutrient concentrations, increased pH, supersaturation of oxygen, and high Coliform bacilli (1800 100ml -1) concentrations have previously been recorded (Grindley & Eagle 1978). It was noted in this study, however, that with the exception of potential pathogenic bacteria, the effects of effluent discharge on water chemistry, substratum, and biota, at the time of sampling, did not appear to be serious (Grindley & Eagle 1978). Later studies indicated elevated nitrogen and phosphorous levels in the region of the sewage works outfall (Grindley & Snow 1983), though due to the short residence time of water in the lower estuary eutrophication is unlikely.

Watling & Watling (1980, 1982) found the concentration of certain metals in sediment and water samples in the estuary to be elevated above true background values as determined for other south-eastern Cape estuaries, though concluded that there was no real indication of metal build up, or of levels which could be considered as a pollution hazard. Calvo-Ugarteburu (1998) measured the levels of metals in the soft tissue of oysters ( Crassostrea gigas ) in 1998 finding that, with the exception of lead and cadmium, levels similar to those found by Watling & Watling (1976). None of these measured levels is above the maximum tolerable standards for heavy metal in the water column in South Africa (DWAF 1995).

Calvo-Ugarteburu (1998) reported DDT and HCB as not being detectable in sediments below oyster culture racks in Knysna Estuary, while Dieldrin and DDO returned levels ranging from 0.01 ng g -1 to 7 ng g -1 dry mass. Fresh oyster tissue collected at the same time gave traces of DDD (3.5-10.6 ng g -1); DDT 0.08 ng g -1 and HCB 0.07-0.60 ng g -1. Dieldrin was not detectable. None of these results is cause for concern at present (Allanson et al . 2000).

management plan GRNP Appendices 15

The development of the Knysna Quays Marina in 1996 exposed sands contaminated with creosote. Investigating the possible presence of polynuclear aromatic hydrocarbons, Allanson et al. (2000) found that napthalene was the dominant moeity in the creosote-contaminated groundwater seepage (3200 g l -1) followed by fluorane (74 g l -1) and phenanthrene (25 g l - 1). The remainder of the array of chemical components commonly found in creosote approached or were below detectable limits (0.1 g l -1). An initial creosote residue (as phenol) of 350 g l -1 was found to decay to close to 40 g l -1 within 24 days once the seepage was contained.

Tsitsikamma systems

Harrison et al . (1995, 1996), using seven parameters (dissolved oxygen, oxygen absorbed, unionised ammonia, E. coli . nitrate nitrogen, phosphate phosphorous and chlorophyll- a) calculated a water quality index rating value (between 0 (poor) - 10 (good)) for the various estuaries along the Cape Province. The index ratings for the estuaries (or river mouths) within the Tsitsikamm region ranged from approximately 5.5 to 9.0, with elevated E. coli levels impairing the water quality of several of these estuaries. Additional water quality data exists for the estuarine section of the Sout / Salt River and Groot River (west) (Morant & Bickerton 1983), as well as the freshwater section of the Salt River (De Moor et al . 2004).

RIVER MANAGEMENT

Many of the challenges SANParks faces with respect to managing river ecosystem in national parks are common to all parks. Prominent issues include:

Fragmented catchment ownership: For most rivers in parks, only portion of river ecosystems or catchment occur within the park boundaries. In TNP only the very lower reaches of the major rivers (Salt, Groot [West], Bloukrans, Lottering, Elandsbos, Storms, Elands, Groot [East]), and the upper reaches of the Palmiet River in the contractually managed Soetkraal properties, occur within the park. Many anthropogenic changes to rivers originate in catchment areas outside of parks, and are consequently processes or activities over which SANParks has little or no influence.

Reduced ecosystem variability: Scientific studies have demonstrated that the maintenance of inherently variable physical processes, and in particular variability in the flow of water, is essential for healthy functioning of river ecosystems. Past river management by government agencies, however, was frequently directed at minimizing fluctuations in flow to ensure stability of supply for off-channel, non-ecological uses. This has resulted in a legacy of, for example, water storage behind dams and regulating flow for irrigation or domestic supply, which presents considerable operational challenges when attempting to achieve a new ideal of managing for healthy river ecosystem rather than just predictable water supply.

Legislated management: Section 3 of the National Water Act (Act 36 of 1998) clearly identifies the National Government as the public trustee of the nation’s water resources, which acting through the Minister of Water Affairs and Forestry has the power to regulate the use,

management plan GRNP Appendices 16 flow and control of all water in South Africa. SANParks thus does not directly, and in most cases also indirectly, manage hydraulic processes and resource use in rivers. The Act also states that the Department of Water Affairs and Forestry must devolve most of the catchment management issues to Catchment Management Agencies (CMA’s) that include representatives of local interest groups and relevant government agencies. Although CMA’s will provide opportunity for cooperative catchment management, the Tsitsikamma-to-Fish CMA to whom management of river catchments in the Tsitsikamma area will be devolved, has not yet been established.

The most productive future role for SANParks in the management of rivers in parks would be the active participation in structures and processes for cooperative catchment management involving all stakeholders. Via such mechanisms the case could be made for resource utilization that is not only equitable and efficient, but also results in the protection of a healthy aquatic environment for present and future generations.

Prominent activities would most likely include lobbying for and active participation in determination of ecological reserves; facilitating assessment of ecosystem and river health; and provision of information and insight obtained through research and monitoring to facilitate informed decision making and the successful implementation of catchment-scale adaptive management systems.

MARINE AND COASTAL PROCESSES

The Tsitsikamma MPA is situated near the centre of the Agulhas Bioregion, which extends 1706 km from the Mbashe River to Cape Point (Lombard et al. 2005). Nineteen percent (or 327 km) of the bioregion’s coastline is fully or partially protected through MPA’s or closed areas (Lombard et al . 2005) with the Garden Route N.P. contributing four percent ((Lombard et al. 2005). The MPA includes approximately 61 km of rocky shoreline and 5 km of sandy beaches, extends between 0.5 and 3 nautical miles offshore and has a surface area of some 340 km². Although the shoreline is primarily rocky, the subtidal areas consist chiefly of soft bottom sediments ( c. 79%) and to a lesser extent dispersed gravel platforms and rocky reefs (c. 21%) (Flemming et al. 1986). The adjacent 9 km long De Vasselot section, which was excluded from the MPA, is open to extractive resource use by the public in accordance with the Marine Living Resources Act (1998).

The marine fauna of the MPA includes a variety of mammals (dolphins, whales & seals), birds (gulls, terns, gannets & penguins), fishes (reef & migratory species) and invertebrate reef fauna (sponges, ascidians & crustaceans). However, the MPA is too small to provide permanent residence for most of the marine mammal and seabird species recorded for the park, but it is large enough to provide an important feeding ground and/or nursery area. Spawning grounds for commercially exploited chokka squid and all life stages of 17 fish species which are commercially and/or recreationally exploited have been recorded in the park (Sauer 1995; Wood et al. 2000).

In the marine environment extreme endemism is relatively rare (Hockey & Branch 1994), and the ecological case for protection of an area is often based on the safeguarding of an important habitat for commercially or recreationally important species, or protection of a particularly good example of a habitat type and its associated communities (Robinson & De Graaff 1994).

management plan GRNP Appendices 17

The biggest threat to the marine environment in this southern Cape region is extractive living marine resource use (Lombard et al. 2005). Therefore, a key conservation function of the Tsitsikamma MPA is to provide a safe and pristine refuge for exploited fish and invertebrate species.

management plan GRNP Appendices 18

APPENDIX G

IMPLEMENTATION PLAN FOR ESTUARY MANAGEMENT: GARDEN ROUTE NATIONAL PARK

INDEX Page

1. Artificial manipulation of estuary mouths

2. Cutting of aquatic plants (interleading channels

& disturbance simulation)

3. Sediment removal in Touw system

4. Wetland Rehabilitation

5. Law enforcement

6. Resource Utilization

7. Boat mooring/carrying capacity

8. Managing pollution inputs

9. References

1. Artificial Manipulation of Estuary Mouths

management plan GRNP Appendices 19

1. 1 Relevant Objectives

• Reduce the probability of inundation of residential areas on adjacent floodplains whilst at the same time maintaining sufficient head of water to maximise the scouring of sediments from the estuary mouth on breaching and maintain natural inundation flux in marginal wetlands, by maintaining the system of artificially breaching estuaries in the Touw, Swartvlei and Groot (West) systems.

• Develop an improved understanding of the hydrodynamics of priority estuarine systems, where possible emphasizing linkages between freshwater, estuarine and marine systems.

1.2 Purpose

Regularly artificial breaching the Touw, Swartvlei and Groot (West) estuaries when the water level is lower than when breaching could occur naturally is in most instances undertaken, in the cases of Touw and Swartvlei, to reduce the threat of flooding of residential and other properties that have been developed on the estuarine floodplains, and in the case of Groot (West) to prevent flooding of some facilities (access road, parking area, septic tanks) in waterfront properties in Natures Valley, and camping sites in the SANParks Nature’s Valley restcamp. Premature breaching and closure of estuaries may at times also be undertaken when considered essential for the restoration of natural patterns and processes and reduction of the effects of pollution.

1.3 Context

SANParks managed conservation areas from Wilderness to Tsitsikamma incorporate several estuarine systems. These include one estuarine bay (Knysna), one permanently open estuary (Salt) four temporarily open estuaries (Touw, Swartvlei, Noetsie, Groot (West)), four estuarine lakes (Swartvlei, Rondevlei, Langvlei, Eilandvlei) and 16 river mouths (Kranshoek, Brak, Helpmekaars, Klip, Bloukrans, Witels, Lottering, Elandsbos, Geelhoutbos, Kleinbos, Storms, Bruglaagte, Langbos, Sanddrif, Elands, Groot (East)). The overwhelming majority of issues associated with manipulating estuary mouth states are in three of the four temporarily open estuaries (Touw, Swartvlei, Groot (West)) which have residential areas and related infrastructure inappropriately located on adjacent floodplains.

Historical records indicate that the Touw, Swartvlei, Groot (West) are temporarily open/closed estuaries. As such these systems are blocked off from the sea for varying lengths of time by a sand bar which forms at the mouth. This occurs during low river flows combined with long-shore sand movements in the near-shore marine environment. In the Groot (West) system available morphological evidence coupled with low salinity in the back-barrier suggests that when open the mouth serves mainly as an overflow channel rather than a tidal inlet (Harrison 1996).

Development of residential properties in the towns of Wilderness (Touw Estuary), Sedgefield (Swartvlei Estuary) and Natures Valley (Groot (West) Estuary) on the estuarine floodplains have in the past been approved by local municipalities, and new developments on floodplains continue to be approved by local governments. If the estuaries are left to breach naturally, water levels under a typical flooding regime can result in inundation some residences in Wilderness in the case of the Touw

management plan GRNP Appendices 20

Estuary, Sedgefield in the case of Swartvlei Estuary, and an access road in Natures Valley; the public parking area near the estuary mouth, private gardens along Lagoon drive, and some camping sites in the SANParks De Vasselot restcamp in the case of the Groot (West) estuary. High water levels in the Groot (West) also result in the flooding of septic tanks associated with some waterfront properties, which in turn results in faecal contamination of the estuary indicated by high E. coli counts, particularly during draw-down periods. Flooding of land adjacent to estuaries can even be expected when river floods coincide with high tides when the estuary is open, especially in the Sedgefield Extension 1 area adjacent to Swartvlei Estuary (Howard-Williams & Allanson 1979).

In the Touw Estuary, expected water levels (meters amsl) that would result during flood events of different return frequency are 5 yr. = 3.22; 10 yr. = 3.61; 20 yr. = 3.88; 50 yr. = 4.01; 100 yr. = 4.93 (Görgens 1979) which similarly would result in the flooding of waterside properties. CSIR (1981, 1982) concluded that maintaining the height of the sand sill at the estuary mouth at between 2.1 m and 2.4 m amsl should prevent flooding of low-lying properties by floods with a return frequency of 1 in 50 years or lower. In the case if Swartvlei Estuary it was recommended by Howard-Williams & Allanson (1979) and Whitfield et al . (1983) based on hydrological modelling undertaken by CSIR (1978) and ecological considerations that artificial breaching should occur at 2.0 m amsl. Hydraulic studies of the Groot Estuary are extremely limited, being confined largely to unpublished once-off measurements of flow speeds by ECRU in 1982 (Morant & Bickerton 1983). Local knowledge suggests that the difference between the water height at which flooding of facilities may commence (2.0m amsl) and estimated average natural breaching height (2.1-2.2m amsl) is relatively small. Higher water levels could be expected during periods of when the sand sill builds up higher than 2.2m amsl, or when river floods coincide with periods when the estuary mouth is closed, or even with high tides when the estuary is open.

The height at which estuaries are artificially breached are in essence a compromise to reduce the threat of flooding of residential and other properties that have been developed on the estuarine floodplains, whilst retaining sufficient head of water on breaching to perpetuate hydrological processes influencing the erosion of marine sediments in the estuary mouth and maintain patterns and processes dependant on varying estuarine water levels.

1.4 Management actions

The following operating principles to the breaching of estuaries in the Garden Route National Park, Wilderness National Park, Tsitsikamma National Park and Knysna Protected Environment apply:

1.4.1 Permissibility of artificial breaching of estuary mouths

Artificial breaching of an estuary by SANParks may be undertaken where:

The natural breaching of an estuary is significantly disrupted by human alteration of hydraulic processes and such change in the breaching is deemed by scientific investigation to have had, or be likely to have a long-term detrimental effect on estuarine biota or environmental processes.

management plan GRNP Appendices 21

Natural breaching will result in the flooding of residential properties adjacent to the estuary that had been developed prior to SANParks implementation of estuary management on proclamation of the estuary as portion of a national park or protected environment.

1.4.2 Sandbar skimming (Touw Estuary, Swartvlei Estuary, Groot (West) Estuary).

During periods when an estuary is closed the height of the sandbar at the estuary mouth may be reduced by use of earth moving equipment operated by, or under the direct supervision of authorised SANParks personnel. Skimming heights and frequency pertaining to estuaries are:

Touw Estuary

A portion of the sandbar at the Touw Estuary mouth (stretching from the estuarine waters edge to the sea high water mark, and a width of not wider than 5 meters will be maintained at a height of between 2.1 and 2.4m amsl.

Measurement of the maximum estuary sandbar height will be undertaken at intervals of not greater than seven (7) days during periods when the estuary mouth is closed.

Every Monday, the height of the sandbar is measured with a dumpy level to obtain the height of the sandbar.

Should the sandbar have reached a height of more than 2.4 amsl, a bulldozer is brought onto the beach skim the level down to not below 2.1 amsl, another measurement is taken to ensure that the desired height is achieved. In the event of high seas and severe weather (rain) warnings, the re- measuring of the sandbar height will be undertaken on any other day.

Swartvlei Estuary

The sandbar at the estuary mouth will not be regularly skimmed in order to maintain it at a fixed height.

Under what conditions will bar be skimmed?

Measurement of the maximum estuary sandbar height will be undertaken at intervals of not greater than seven (7) days during periods when the estuary mouth is closed, and will be recorded within the monthly report

Where skimming is undertaken the sandbar height may not be reduced to less than 2.0m amsl over any portion of its length.

Groot (West) Estuary

The sandbar at the estuary mouth will not be regularly skimmed in order to maintain it at a fixed height.

Under what conditions will bar be skimmed?

• Excessively high levels of sand deposits at the mouth which would not allow for natural breaching.

management plan GRNP Appendices 22

Where skimming is undertaken the sandbar height may not be reduced to less than 2.0m amsl over any portion of its length.

A log to be maintained of natural and artificial breaching. Sandbar skimming will not be undertaken at any estuary other than Touw, Swartvlei and Groot (West).

1.4.3 Preparation for breaching of estuaries

A preparatory channel may be constructed at any time prior to estuary breaching if deemed necessary by SANParks personnel to facilitate future breaching. To be proactive regarding possible flooding, a preparatory channel will be prepared when the water level reaches 1.8 amsl (Touw Estuary) and 1.6 amsl (Swartvlei Estuary) within the months of March to April and August to December. The extent and design of a preparatory channel must be such that premature breaching of an estuary (<2m amsl – Swartvlei Estuary; < 2.1m amsl – Touw Estuary, <2.0m amsl – Groot (West) Estuary) cannot occur as a result of either sediment erosion from the preparatory channel by high seas, or deliberate acts to breach the estuary by unauthorised persons. The widths of the channels should not exceed 5m for the Touw Estuary and 10m for the Swartvlei Estuary.

The Grootriver Estuary preparatory channel would be undertaken as follows: A preparatory channel is constructed by means of a Bulldozer, &/or Front-end Loader, in an area that has been identified where a natural breach is most likely to take place, from the waters edge of the estuary to the high water mark on the seaward side, and the height of the plug on the seaward side not exceeding 3 amsl and not lower than 2 amsl. The channel is skimmed to a level (or below) where natural flow would take place. The sand removed from the channel, should either be distributed adjacent to the channel on the beach or stockpiled and removed at a later stage in accordance to the resource utilization policy .

1.4.4 Estuary Breaching (Touw Estuary, Swartvlei Estuary, Groot (West) Estuary).

The breaching of estuaries will be undertaken solely by, or under the direct supervision of authorised SANParks personnel. In the case of the Groot (West) Estuary breaching, may at times be undertaken jointly by Bitou Municipality and SANParks.

Wherever possible it should be attempted to coincide estuary breaching with a receding tide to maximise the rate of sediment erosion in the estuary mouth by out-flowing water.

Wherever possible heavy earth moving equipment, such as a bulldozer, mechanical shovel, or similar such equipment should be used in the breaching of an estuary to maximise the length and depth of the breaching channel; increase the probability of being able to effect a breach at the desired time; and decrease the time required to effect a breach during periods of increased flooding risk. Environmental authorisation for the method of breaching (i.e. bull dozers) the Touw and Swartvlei estuaries has been obtained from the Department of Environmental Affairs.

Breaching of the sandbar at an estuary mouth to effect a linkage between the estuarine and marine environment will, except where conditions listed under 4.5 below apply, in (a) Swartvlei

management plan GRNP Appendices 23

Estuary be undertaken when the water level within the estuary attains a height of not less than 2.0m amsl as measured at the most seaward portion of the estuary mouth, (b) in Touw Estuary be undertaken when the water level within the estuary attains a height of not less than 2.1m amsl and not more than 2.4m amsl as measured from the gauge plates in both the Touw and Swartvlei Estuaries and (c) in the Groot (West) Estuary be determined by authorised SANParks personnel on a cases by case basis in conjunction with the Bitou Municipality but will not be when the water level in the estuary is less than 2.0m amsl as measured at the most seaward portion of the estuary mouth.

Wherever possible it should be attempted to coincide estuary breaching with a receding tide to maximise the rate of sediment erosion in the estuary mouth by out-flowing water.

Where ever possible the estuary should be breached 3-4 days before springtide.

Wherever possible heavy earth moving equipment, such as a bulldozer, mechanical shovel, or similar such equipment should be used in the breaching of an estuary to maximise the length and depth of the breaching channel; increase the probability of being able to effect a breach at the desired time; and decrease the time required to effect a breach during periods of increased flooding risk.

It is preferable to be breach the estuaries during September to November as this corresponds to breeding cycle of certain aquatic species.

• It is recommended that breaching is undertaken during a receding tide to maximise the rate of sediment erosion.

• The current methodology will be regularly assessed and adjusted to implement the most effective breaching method.

• Law Enforcement personnel to be in attendance during the opening of the estuary mouth to assist with crowd control & to prevent fishing during the initial mouth opening.

1.4.5 Premature breaching of estuaries

Breaching of the sandbar at an estuary mouth to effect a linkage between the estuarine and marine environment where the water level in the estuary is, in the case of the Touw Estuary below 2.1m amsl, and in the case of Swartvlei and Groot (West) estuaries is below 2.0m amsl, can be undertaken if one or more of the following conditions occur:

Prevailing meteorological conditions or forecasts as provided by the Eden Disaster Management (or South African Weather Services in the case of Grootriver) will assist with the decision for premature breaching of the estuaries. If 50mm rain is predicted and actually falls breach @ 1.8, 100mm predicted breach @ 1.6, 150mm predicted breach @ 1.4

A log is to be kept of all information received from disaster management (time, date, extent of predicted rainfall, estuary height at time, local rainfall measurement, actions undertaken ).

The undertaking of emergency repair of essential services infrastructure.

management plan GRNP Appendices 24

The health of public who may come into contact with the estuarine water is at risk, as indicated by accepted virological, bacteriological or chemical tests, and/or where there is the obvious introduction of matter that may pose a significant health risk as determined by SANParks personnel and where necessary as advised by an appropriate qualified and competent authority.

Where the opportunity to prevent exposure of the public to physical contact with the water does not exist, and breaching of the estuary mouth can be demonstrated to have a high probability of significantly reducing the health risk.

Facilitation of the dispersal, dilution or removal of chemical pollutants that have been accidentally introduced into the system, and which pose a significant threat to either public health, the survival of estuarine biota, or the ecological functioning of the estuarine system.

There has been no time in a 24 month period where the estuary has remained continually open for one complete tidal cycle (28 days) or longer, and where such an extended closed period is indicated by scientific investigation to have, or likely to have, a long-term detrimental effect on estuarine biota, or result in salinity levels exceeding the range which would normally occur in the estuary.

There is scientific evidence that the failure to open the estuary would result in significant direct or indirect threat to red-data and/or endemic species, and where the breaching of the estuary would significantly reduce such threat.

Significant change occurs in the distribution and abundance of estuarine biota, or substantial die- off of biota occurs, and where such changes will have a significant effect on estuarine biota or the ecological functioning of the estuary or adjacent marine ecosystem, and can be directly attributed to the estuary being closed.

1.4.6 Premature closure of estuaries (Touw, Swarvlei, Noetsie, Salt, Groot (West) estuaries)

An estuary or river mouth may not be artificially closed other than to prevent the inflow of pollutants from the marine environment.

1.4.7 Unmanipulated estuaries

Artificial manipulation of the mouth state (both breaching and closure) will not be undertaken or attempted at the Knysna estuary and the Kranshoek, Brak, Helpmekaars, Klip, Bloukrans, Witels, Lottering, Elandsbos, Geelhoutbos, Kleinbos, Storms, Bruglaagte, Langbos, Sanddrif, Elands, and Groot (East) river mouths.

1.4.8 Estuarine modelling

Investigations will be made into securing funding for the undertaking of hydrodynamic modelling of the Swartvlei system. The primary objective of such a modelling exercise would be to provide guidelines to manage the hydrology of the estuary by achieving a balance between ecological

management plan GRNP Appendices 25 considerations and the risk of flooding. The proposed modelling would incorporate more advanced modelling techniques than have been used in past assessments including improved understanding of linkages between freshwater, estuarine and marine systems, and consideration of sedimentary processes, and would be based on more current bathymetric assessments of the estuary systems and topography of its immediate surrounds.

2. Cutting of aquatic plants (interleading channels & disturbance simulation) in the Wilderness lake systems

2.1. Relevant Objectives

• Increase connectivity between waterbodies in the Touw system by regular cutting of emergent and submerged aquatic plants in the Serpentine channel, and interleading channels between Eilandvlei, Langvlei and Rondevlei.

• Management of aquatic plants in the Touw system to reduce the trend towards single species domination by manipulating flooding and/or cutting of Phragmites stands to simulate natural disturbance at selected sites in Serpentine, Eilandvlei, Langvlei and/or Rondevlei.

2.2 Purpose

Cutting of submerged and emergent aquatic plants in channels within the Touw system may be undertaken to maintain connectivity between waterbodies to facilitate water movement and maintain opportunities for the movement of aquatic biota between waterbodies. Restricted cutting of emergent aquatic plants may also be undertaken in front of bird-hides in the Wilderness lakes to enable better viewing of waterbodies from these facilities, and submerged aquatic plants in selected boat slipways in Swartvlei and Eilandvlei to facilitate boat launching. C utting of emergent plants may be also be employed by SANParks for the purpose of altering plant distribution and establishment pattern where the objective of such management is to reinstate appropriate biodiversity patterns and processes.

2.3. Context

The Wilderness lakes are characterised by supporting extensive stands of both submerged and emergent aquatic plants (macrophytes). Abundant submerged species in lakes include Potamogeton pectinatus (fennel-leaved pondweed) , Ceratophyllum demersum (hornwort) and Charophyta (stoneworts), whereas Zostera capensis (eelgrass) and Ruppia cirrhosa (spiral tasselweed) occur in Swartvlei Estuary. Widespread and abundant emergent species include Phragmites australis (common reed), Typha capensis (bulrush) and Schoenoplectus scirpoideus (no common name) (Weisser & Howard-Williams 1982; Russell 2003).

management plan GRNP Appendices 26

Macrophyte growth in the interleading channels in the Touw system is considered to be a major factor inhibiting water flow between lakes (Hall 1985a). The movement of fishes between waterbodies may also be inhibited by extensive macrophyte growth in channels. Fish communities are typically dominated by juvenile marine species (Hall 1985a, 1985b; Kok & Whitfield 1986; Hall et al . 1987; Russell 1996); with the number of species declining the further removed a waterbody is from the sea (Hall 1985a; Russell 1996). This suggests that several fish species may experience difficulty in negotiating the inter-connecting channels between lakes (Hall et al . 1987), which would be exacerbated by abundant plant growth. To facilitate movement of water between lakes, and maintain opportunities for migration of marine fishes, aquatic plants in interleading channels in the Touw system are periodically cut by SANParks to a depth of approximately 1 m below water level using a mechanical reed cutter.

Howard-Williams (1979) provided recommendations for the cutting of aquatic plants in the Swartvlei system, emphasizing the need for controlling aquatic plants only in areas where there is direct and unavoidable conflict with human use of the waterbody. In some cases where the occurrence of aquatic plants has hampered human recreational use of the Wilderness lakes, the offending stands have been cut or removed, at times with the permission of SANParks, but in many instances illegally. For example, many owners of residential dwellings adjacent to the lower portions of the Touw Estuary remove shoreline vegetation, including Phragmites australis stands, either to improve their view of the waterbody, or to construct jetties, slipways or retaining walls (Russell 2003). Furthermore, privately owned holiday resorts on the western bank of Swartvlei Lake maintain access channels to jetties by cutting submerged aquatic plants, and shoreline vegetation is removed presumably to facilitate access to the water edge (Russell 2003).

Long-term changes have been recorded in the distribution and abundance of emergent aquatic plant communities in the Touw system, chacterised by and increase in the distribution of Phragmites australis (53.9ha; +53%), Grass or fields (23.1ha; +35%) and Scrub or trees (12.2ha; +45%). Substantial declines occurred in the distribution of Juncus kraussii (76.2ha; –243%), Schoenoplectus scirpoideus (10.1ha; –38%) and Low scrub or fynbos (7.8ha; –66%) (Russell 2003). Probable causes of change in the distribution of wetland plants include the natural tendency of plants to colonise new areas, as well as anthropogenic manipulation of physical, chemical and biological processes, including withdrawal of disturbance (Russell 2003). Potential management control methods include the flooding of wetland areas in association with mechanical disturbance (Russell & Kraaij 2008).

2.4. Management actions

2.4.1 Circumstances where cutting of aquatic plants is permitted

Channels in Touw System: The cutting of emergent and submerged aquatic plants will be undertaken in areas of the Serpentine channel, the channel between Rondevlei and Langvlei, and the channel between Langvlei and Rondevlei where plants are sufficiently numerous and widespread to significantly impede water flow within the channels. Plants cut will include the above ground portions of emergent plants which intrude into the channels, and both emergent plants and submerged

management plan GRNP Appendices 27

aquatic plants rooted within channels. Plants or plant portions within channels will be cut to a level of approximately one meter below water level using a mechanical plant cutter. The interval between cuttings may vary, depending on the rate of plant regrowth following cutting and the rate of establishment of new stands, but will be at intervals appropriate for enabling continuous free movement of water between waterbodies. The cut portions plants will be removed from the waters. Cut plant material may be deposited on the channel banks at a height where it will not be re-suspended by annual flooding (>3m amsl).

Rondevlei and Malachite bird-hides: Cutting of emergent aquatic plants can be undertaken in front of the Rondevlei and Malachite bird-hides. In the case of the Rondevlei hide which during low water levels is set back from the waters edge, the area cut may not exceed the width of the hide at the front of the hide, and twice the width of the hide at the waters edge. In the case of the Malachite hide which is positioned in an area which is permanently inundated, the width of the area cut may not exceed twice the width of the hide. Aquatic plants which overhang boardwalks by which the bird-hides are accessed will be periodically cut to facilitate use of these facilities.

Boat launching: Cutting of submerged aquatic plants to facilitate boat launching is permissible in selected localities. These include all slipways utilised by SANParks for access to waterbodies for management or research, and the public launch facility on the western shore of Eilandvlei. Cutting of aquatic plants available for public access to Swartvlei Lake and Eilandvlei and will be considered on a case by case basis. Cutting will be restricted to no more than two public accessible slipways per waterbody. Cutting of plants in front of such slipways will only be undertaken either by, or under the direct supervision of authorised SANParks personnel. Cutting of aquatic plants in front of private slipways or jetties will not be permitted.

Management of plant species encroachment: Long-term monitoring has demonstrated changes in the distribution and abundance of emergent aquatic plants in the Touw system, with Phragmites reed becoming more widespread and abundant. Management intervention to halt or reverse this trend may be considered desirable, and physical removal of selected species in target areas may be considered to be a preferred management technique. Cutting of emergent aquatic and other wetland plants by SANParks is permissible in any area where it is deemed appropriate for the purpose of manipulating plant distribution and establishment pattern where the objective of such management is to conserve biodiversity patterns and processes.

2.4.2 Circumstances where cutting of aquatic plants will not be undertaken or permitted

Eroding banks: Cutting of emergent aquatic plants will not be undertaken, and will not be permitted in areas where banks are eroding or at risk to erosion, as a result of periodic high flow speeds, natural wave action, or waves resulting from usage of waterbodies by powerboats. These areas include all estuaries and any other area in the estuarine lakes and river mouths where the conditions regarding erosion risk apply.

management plan GRNP Appendices 28

Property views: Cutting of emergent aquatic plants to improve view of waterbodies in front of private waterfront properties including residential properties, smallholdings, farms or holiday resorts will not be undertaken or permitted.

• External systems: Regular cutting of aquatic plants will not be undertaken by SANParks in aquatic systems or portions of systems outside of the National Park or the Knysna Estuary. Permission by external agencies to undertake cutting outside of the protected area will be considered on a case by case basis, where it can be demonstrated that there is a clear benefit to the environment.

3. Sediment removal in Touw system

3.1. Relevant Objectives

• To increase connectivity between waterbodies in the Touw system by removing sediment which has accumulated at point localities in interleading channels due to anthropogenic factors

3.2. Purpose • Eliminate artificially created restrictions to flow in the Touw system

3.3 Context • Interleading lakes • Flow exchange via interleading channels • Surrounding development with channelized runoff • Bridges over channels, with deposition over the sides • Resulting in restrictions to flow

3.4 Sediment removal in other estuary systems

Sediment removal in other estuarine systems will only be considered on a case by case basis where it can be demonstrated from formal surveys that ongoing and continued accumulation of sediments is occurring within the system and which is having a negative effect on physical and biological patterns and processes, and where sediment removal can be demonstrated to have a high probability of significantly reducing this negative effect .

4. Wetland Rehabilitation

4.1. Relevant objectives:

• Rehabilitation of estuarine wetland areas with emphasis on the reestablishment of indigenous vegetation in intertidal and supra-tidal areas where changes have resulted from past landuse practices, system manipulation and extralimital invasion. • To rehabilitate wetland systems in the GRNP • To control accelerated erosion in the GRNP resulting from human activities

management plan GRNP Appendices 29

4.2. Purpose:

The purpose of wetland rehabilitation programs in the GRNP could include one or more of the following: • Reinstate natural topography. • Elimination of alien plants. • Disposal of dumped material. • Encourage of re-growth of indigenous vegetation. • Erosion prevention • Reinstate natural wetland processes • Raise awareness of the importance and function of wetlands.

4.3. Context:

Due to the proximity of wetlands in the Garden Route National Park to towns and their associated infrastructure, wetlands are continuously under threat. The fragile nature and importance of wetlands is often not properly understood and the disturbance and infill thereof was historically not seen as problematic. In a number of cases this sentiment still exists today. It is of crucial importance that wetlands within the Garden Route National Park are not disturbed by, or lost to, any form of unnatural pressures. Furthermore concerted efforts must be made to ensure that wherever possible disturbed wetland areas are rehabilitated.

4.4. Management Actions:

It is foreseen that rehabilitation programmes could have positive environmental outcomes in the following instances:

• Re-establishment of indigenous vegetation where disturbance has taken place on estuary margins for e.g. infrastructure and other developments. • Removal of defunct structures, jetty’s, slipways, bridges etc. • Re-establishment of indigenous vegetation in inter-tidal and supra-tidal areas where changes have resulted from past land use practices, for example the disused airfield on George Rex Drive on the Knysna estuary. • Re-establishment of indigenous vegetation in areas where oyster racks have been removed in the Knysna estuary. • Re-instatement of natural processes in influent rivers in the Knysna system to reduce sedimentation and pollution e.g. proposed Working for Wetland projects in the Bigai and Bongani streams. • Rehabilitation of estuarine wetland areas with emphasis on the system manipulation and extralimital invasion.

Specific management actions will be defined on project initiation and dependent of the availability of resources.

Where remedial actions are undertaken by external agencies, SANParks will operate in an advisory role during planning and implementation phases.

5. Law enforcement (relevant to resource usage)

management plan GRNP Appendices 30

Law enforcement pertaining to estuarine management is discussed in more detail in the relevant safety and security plans of the various management units within the Garden Route National Park. Below are the main management actions to ensure compliance and enforcement of legislation applicable to estuarine management and the maintenance of ecological integrity of the estuaries within the GRNP.

5.1 Management actions

• Enforce legislation pertaining to activities that impact on estuarine ecosystems and their functioning; • Enforce, through compliance monitoring, existing legislation under the MLRA e.g. permits, catch restrictions, use of cast nets. • Eliminate illegal activities (eg. gill netting); • Prohibit capture of fish during mouth breaching events in temporarily open/closed systems • Prohibit capture of fish with seine nets, gill nets and traps; • Restrict collection of mudprawn, sand prawn, blood worm and pencil bait only using legal implements; • Eliminate the collection of any bait organism within the designated invertebrate reserves; • Promote alternatives to consumptive exploitation eg. Catch and release fisheries, eco-tourism; • Identify protected areas for the conservation of over-exploited linefish species (eg. Dusky cob & white steenbras). The area must include the mouth and adjacent marine environment; • Prioritise and increase funding for enforcement and research;

management plan GRNP Appendices 31

6. Resource Use

6.1 Relevant Program Objectives

• Monitor marine biota, including resource stocks, harvesting, and alien invasions. • Quantify current extractive resource activities. • Quantify attitude of fishers towards compliance & MPAs

6.2 Purpose

Recreational and subsistence fishing within the Garden Route National Park currently takes place within the De Vasselot portion of the Tsitsikamma Section, the Knysna Estuary, the Swartvlei system (estuary and lake) and within the Touw estuary and Island Lake of the Wilderness Lake system in accordance with the regulations of the Marine Living Resources Act (1998).

The current monitoring programme aims to assess the recreational and subsistence fisheries occurring within the above estuarine systems and hence managed by SANParks. Resource monitoring provides information on trends in participation and exploitation of fishery and bait resources within different systems / areas and the findings provides much needed information that can feed directly into management decisions and aid in maximising the long-term economic benefits at both a local level and a national level by conserving stocks of important linefish species.

Relevant guideline actions (info only, not for direct inclusion unless part of purpose)

• Identify and quantify (monitor) consumptive resources, their value, and the present levels of exploitation in all systems • Reduce the number of fishing competitions • Reduce bag limits on all threatened estuarine angling species • Research & monitoring

6.3 Context

Recreational angling is a popular and growing activity in South Africa. In 1995 the estimated number of participants within this fishery was around 412 000 with an annual increase of around 2% (McGrath et al . 1997). The smaller subsistence sector was estimated to consist of around 29 000 individuals with the majority being found on the East coast (Clarke et al . 2002). Subsistence fishers harvest a variety of species from invertebrate bait and food organisms to linefish species, which they rely on as a source of protein or to sell (Clarke et al . 2002, Branch et al . 2002). The collective annual harvest by all sectors was estimated at 4.5 million fish weighing 3000 tonnes each year (Brouwer et al . 1997). Estuarine fishing has historically been a popular activity for both coastal residents and visitors, being particularly active areas for both recreational and subsistence anglers due to the abundance of fish, the ease of accessibility and the protected waters, (Baird et al. 1996). Trends indicate that estuarine angling has increased in popularity, particularly since the ban on beach driving (MacKenzie 2005), with an estimated 70 000 participants (Lamberth and Turpie 2003). management plan GRNP Appendices 32

Despite the fishery management strategies employed by government (size restriction, bag limits, closed seasons) and amendments implemented over time, various studies suggest that catch rates in recreational fishing along the South African coastline have decreased and species composition of catches have changed (Crawford & Crous 1982, Hecht & Tilney 1989, Brouwer et al . 1997, Penney et al . 1999, Griffiths 2000, Brouwer & Buxton 2002). Based on trends in catch-per-unit-effort, research has shown that stocks of most of South Africa’s linefish species are currently over-exploited (Griffiths 2000).

Extractive over-exploitation of resources has been highlighted as a both a threat to estuarine ecosystems (Whitfield 1997) and the greatest threat to marine biodiversity (Lombard et al. 2005). In particular estuaries are highly productive acting as both nursery and feeding areas for a number of important linefish species, several of which form important components of the recreational and commercial linefisheries. Careful management is required in order to ensure adequate protection of resources and maintaining the sustainability of the linfishery. Consequently, it is essential that long term monitoring programmes to evaluate catch and effort and the sustainability of fishing and bait collecting occurring within the estuarine systems of the GRNP be implemented.

6.4 Management Action

Monitoring will follow a randomly stratified sampling design with equal monthly effort within each estuary body. The program consists of monthly direct-contact roving creel surveys and instantaneous user counts undertaken during daylight hours (06:00 – 18:00) – representing daylight fishing effort only. On each survey day two instantaneous counts covering the entire survey area are completed to provide information on spatio-temporal distribution of fishing/ bait collecting effort.

During the roving creel surveys, only those people actively involved in resource utilisation (bait collecting or fishing) or packing-up after their outing are interviewed. Two questionnaires are used. A sub-sample of the anglers are asked a comprehensive questionnaire used to gather information such as: (i) user demographics; (ii) angler attitudes and perceptions towards management; (iii) fishing expenditures; (iv) species composition and size distribution of catches made; (v) effort (duration of fishing trip, number of lines in use); and (v) bait dynamics. A second questionnaire, used the majority of the time and on repeat encounters with anglers, is aimed at primarily gathering basic demographics and catch and effort data. Bait collectors are interviewed with similar questionnaires.

6.4.1 Monitoring of bait populations in Swartvlei estuary

management plan GRNP Appendices 33

The impact of bait collecting on the population density of sand prawn (Callianassa kraussi ), the dominant bait organism in the Swartvlei estuary will be assessed through annual burrow counts in three exploited areas and three ‘unexploited / lightly exploited areas. At each of these areas three transect will be done from the low water mark to high water mark, with at least triplicate counts being done at the bottom, middle and top of the main prawn bed. The density of sand prawn burrows in exploited areas will be compared with that of ‘unexploited’ sites to evaluate the extent of decline in prawn densities due to harvesting .

7. Boat launching sites and moorings

7.1 Relevant Program objective :

• To ensure compliance with environmental legislation for all park infrastructure, development and activities

7.2 Purpose

• Reduce traffic on wetlands; • Reduce vegetation destruction on intertidal areas; • Restrict boating activities to appropriate localities; • Eliminate mooring of vessels in high intensity use areas.

7.3 Context

Due to the open nature and accessibility to most areas along the banks of estuary these areas can generally be accessed with a variety of smaller vessels or craft. It is possible for fishers, bait collectors, canoeists, windsurfers and kite-boarders to access the water over sensitive estuarine areas i.e. wetland areas. Vessels are also often landed on inter-tidal wetland areas for the purpose of mooring (or tying up). These are mostly informal moorings which are often temporary/seasonal of nature, however potential damage to the inter tidal areas, and shoreline is always a concern. It is important that all moorings are properly placed, maintained and regulated to ensure proper control of this activity.

7.4 Management actions

• Restrict the launching of craft by means of vehicles to designated slipways; • Encourage the launching of all small craft be undertaken at designated slipways; • Ensure registration of slipways; • Encourage reduction in the practise of beaching of vessels on intertidal wetlands; • Restrict mooring points to designated areas; • Develop and maintain records of all mooring points .

8. Pollution inputs (incl E. coli )

8.1 Relevant program objective: management plan GRNP Appendices 34

• Reduce pollution inputs, including sewage, and unnaturally elevated suspended sediment levels in freshwater systems.

8.2 Purpose

• Control pollution and waste emissions including sewage, detergents, agricultural runoff, urban runoff, industrial waste, solid waste and fossil fuels • Control erosion and runoff associated with infrastructure development (roads, residential and industrial) • Identify and mitigate against impacts resulting from industrial and urban development

8.3 Context

Persistent problems are mostly related to Knysna estuary and a number of other primary drainage systems. The existing problem is mostly related to sewage inflow, and other forms of urban, industrial and agricultural waste/ pollutants. Furthermore the problem is aggravated by outdated infrastructure, infrastructure decay, system breakdown and failure.

Sediment inputs from development activities and incorrect farming practices remain a concern within most catchment areas

8.4 Management actions

• Identify occurrence and extent of pollution problems • Local and regional authorities undertake water quality monitoring within estuaries and also rivers and inlets flowing into estuaries. • Notification when levels exceeded at bathing areas • Assessment of turbidity levels in inflow pipes into Knysna estuary during storm events • Interact with local authorities and land owners • E. coli levels in Storms, Groot & Salt systems • Problems experienced in other areas dealt with on an ad hoc basis • Motivate for the building of the sewerage treatment plant in Nature’s Valley

REFERENCES

Allanson, B.R. & Whitfield, A.K. 1983. The Limnology of the Touw River Floodplain. South African National Scientific Programs Report No. 79. CSIR, Pretoria. 35pp. Allanson, B.R. 1981. The coastal lakes of Southern Africa. In: Day, J.H. (ed) Estuarine Ecology with particular reference to southern Africa. Chapter 15. A.A. Balkema, Cape Town . management plan GRNP Appendices 35

CSIR 1978. Hidrouliese studie van die Swartvlei Estuarium. CSIR Report C/SEA 7805/1. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch. CSIR 1981. Wilderness Report No. 1. Evaluation of prototype data and the application of a numerical model to the Wilderness lakes and Touws River floodplain. CSIR Report C/SEA 8113. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch. CSIR 1982. Wilderness Report No. 2. Evaluation of prototype flood conditions and application of the numerical model to conditions when the estuary mouth was opened . CSIR Report C/SEA 8255. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch. Day, J.H. 1981. Coastal hydrodynamics, sediment transport and inlet stability. In : Day, J.H. (Ed.) Estuarine Ecology with particular reference to southern Africa . A.A. Balkema Cape Town. 7-25. Fijen, A.P.M. & Kapp, J.F. 1995a. Wilderness, Swartvlei and Groenvlei Lakes catchment, water management strategy. Introduction and orientation. Department of Water Affairs and Forestry, Pretoria. 16pp Fijen, A.P.M. & Kapp, J.F. 1995b. Wilderness, Swartvlei and Groenvlei Lakes catchment, water management strategy. Proposed water management strategy, objectives and goals. Department of Water Affairs and Forestry, Pretoria. 40pp Fijen, A.P.M. & Kapp, J.F. 1995c. Wilderness lakes catchment, Touw and Duiwe Rivers, water management strategy. Volume 1: Present situation. Department of Water Affairs and Forestry, Pretoria. 115pp Fijen, A.P.M. & Kapp, J.F. 1995d. Swartvlei Lake catchment, Diep, Klein-Wolwe, Hoëkraal and Karatara Rivers, water management strategy. Volume 1: Present situation.. Department of Water Affairs and Forestry, Pretoria. 114pp Fijen, A.P.M. & Van Zyl, F. 1995a. Wilderness lakes catchment, Touw and Duiwe Rivers, water management strategy. Data report. Department of Water Affairs and Forestry, Pretoria. 100pp Fijen, A.P.M. & Van Zyl, F. 1995b. Swartvlei Lake catchment, Diep, Klein-Wolwe, Hoëkraal and Karatara Rivers, water management strategy. Data report. Department of Water Affairs and Forestry, Pretoria. 50pp Görgens, A.H.M. 1979. Estimated flood hydrographs for certain Wilderness streams. Special Report 1/79. In: Hughes, D.A. & Görgens, A.H.M. 1983. Hydrological Investigations and research in the southern Cape coastal lakes region 1979 -1983: Summary and guide to reports. Department of Geography, Hydrological Research Unit, Rhodes University, Grahamstown. Hall, C.M. 1985a. Some aspects of the ecological structure of a segmented barrier lagoon system with particular reference to the distribution of fishes . Unpublished M.Sc. Thesis, Rhodes University, Grahamstown. Hall, C.M. 1985b. The limnology of the Touw River floodplain Part II. Aspects of the ecological structure subject to floods, drought and human interference. Rhodes University Institute for Freshwater Studies. Investigational Report No. 85/1. 137pp. Hall, C.M., Whitfield, A.K. & Allanson, B.R. 1987. Recruitment, diversity and the influence of constrictions on the distribution of fishes in the Wilderness lakes system, South Africa. South African Journal of Zoology 22 (2): 163-169. Harrison, T.D., Cooper, J.A.G., Ramm, A.E.L. & Singh, R.A. 1996. Health of South African estuaries, Groot (Wes) – Great Fish. Executive Report, Coastal and Catchment Environmental Programme, CSIR, Durban.

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Howard-Williams, C. 1979. Recommendations for aquatic plant cutting in Swartvlei. Rhodes University Institute for Freshwater Studies special report No. 79/1, Howard-Williams, C. & Allanson, B.R. 1979. The ecology of Swartvlei: Research for planning and future management. Water Research Commission, Pretoria. 26pp. Jacot-Guillarmod, A. & Allanson, B.R. 1981a. The Touw River floodplain. Part I. Summary and recommendations. Institute for Freshwater Studies, Rhodes University, Grahamstown Jacot-Guillarmod, A. & Allanson, B.R. (Eds.) 1981b. The Touw River floodplain Part II Hydrology and Hydraulics. Institute for Freshwater Studies, Rhodes University, Grahamstown. 36pp. Jacot-Guillarmod, A. & Allanson, B.R. (Eds.) 1981c. The Touw River floodplain Part III The chemical and biological environment and the impact of man. Institute for Freshwater Studies, Rhodes University, Grahamstown. 137pp. Kok, H.M. & Whitfield, A.K. 1986. The influence of open and closed mouth phases on the marine fish fauna of the Swartvlei estuary. South African Journal of Zoology 21 (4): 309-315. Morant, P.D. & Bickerton, I.B. 1983. Estuaries of the Cape, Part II Synopses of available information on individual systems. Report No. 19 Groot (Wes) (CMS 23) and Sout (CMS 22). CSIR Research Report 418. CSIR, Stellenbosch. 54pp. Russell, I.A. 1996. Fish abundance in the Wilderness and Swartvlei Lake systems: changes relative to environmental factors. South African Journal of Zoology 31(1) : 1-9. Russell, I.A. 1999a. Changes in the water quality of the Wilderness and Swartvlei Lake systems, South Africa. Koedoe 42(1) : 57-72. Russell, I.A. 1999b. Freshwater fish of the Wilderness National Park. Koedoe 42(1) : 73-78. Russell, I.A. 2003. Long-term changes in the distribution of emergent aquatic plants in a brackish South African estuarine-lake system. African Journal of Aquatic Science . 28(2) : 103-122. Russell IA & Kraaij T (2008) Effects of cutting Phragmites australis along an inundation gradient, with implications for managing reed encroachment in a South African estuarine lake system. Wetlands Ecology and Management 16(5) 383-393 . Weisser, P.J. & Howard-Williams, C. 1982. The vegetation of the Wilderness lakes system and the macrophyte encroachment problem. Bontebok 2: 19-40. Whitfield, A.K. 2000. Available scientific information on individual South African estuarine systems. Water Research Commission Report No. 577/3/00. WRC, Pretoria. 217pp. Whitfield, A.K., Allanson, B.R. & Heinecken, T.J.E. 1983. Estuaries of the Cape, Report No. 22: Swartvlei (CMS 11). CSIR, Stellenbosch. 62pp.

management plan GRNP Appendices 37

APPENDIX H

FIRE MANAGEMENT SYSTEM FOR THE TSITSIKAMMA MOUNTAIN CATCHMENT AREA

Compiled by: W.J. Vermeulen Assisted by: C.J. Van der Merwe, A.H.W. Seydack, J. Baard , H. Cunningham, T. Kraaij, (SANParks), D. Nortje, B. Bothma, C. Carstens , M. Davids, T. Kok K. Lourens, T. Notley, T. Scholtz, C. Smit, , (MTO Forestry), W. Erlank & J. Thanyan (EC Parks)

November 2009

INTRODUCTION

A large part of the Tsitsikamma Mountain forms part of the Garden Route National Park. This comprises of State forest land and private land managed as a contractual park area.

The primary management objective is biodiversity conservation, within the constraints of measures required for fire protection. Fire management is an important aspect of catchment management, both for biodiversity/fynbos conservation and fire protection. Although the fire management system presented here was developed in close cooperation with important landowners such as MTO Forestry and Eastern Cape Parks, it is still subject to wider stakeholder consultation.

DESCRIPTION OF AREA

The following information on the physical and biological environment of the Tsitsikamma catchment area was largely drawn from Southwood & De Lange (1984).

Physical environment

The larger Tsitsikamma catchment area includes the Langkloof, Tsitsikamma and Kareedouw mountains. Rivers draining the southern slopes of the mountain range are relatively short and flow through deep gorges, incised through the coastal plain (e.g. Bloukrans River, Groot River, Storms River) on their way to the sea.

The catchment area is underlain by rocks of the Ordovician and Silurian age, collectively referred to as the Table Mountain Group, and forms part of the Cape Super Group. Lithologically it consists of various quarts sandstones with bands of shale. The broad east- west trend of the mountain range and the steep southward dip of the formations are due to folding which occurred during the Cape orogeny. Soils are moderately deep, with rocky, loamy sands overlying hard rock or saprolite (lithosols). Soils are generally acid, leached and with a low base saturation.

management plan GRNP Appendices 38

The Tsitsikamma falls within a transitional zone between the subtropical and temperate regions. Rainfall occurs throughout the year with a slight increase in autumn and spring, and with a mean annual rainfall of about 1150 mm. A marked feature of the Tsitsikamma rainfall is the strong orographic control. The mountain range is a barrier to the moisture-laden coastal winds, causing uplift and precipitation. Rainfall, therefore, may exceed 1000 mm on the high peaks, decreasing to 400 mm on northern slopes and foothills.

Maximum temperatures range between 26 °C in January and 19°C in July, and minimum between 15°C in January to 7°C in July. Warm temperatures, occasionally raising above 38°C, are due to hot, dry bergwinds. The highest mean monthly maximum temperature is recorded in February, with July having the lowest mean monthly minimum and maximum temperatures.

Summer winds are mainly south-east, while in winter dry north to north-west winds predominate. South-west winds occur throughout the year, usually accompanied by rain. Bergwind is common in the region from April to September, periodically raising winter temperatures. Thunderstorms are relatively uncommon.

Biological environment

The largest part of the Tsitsikamma Catchment area comprises of Tsitsikamma Sandstone Fynbos, with smaller areas of Eastern Coastal Shale Band vegetation (Mucina & Rutherford 2006), as well as small pockets of Afrotemperate Forest.

Tsitsikamma Sandstone Fynbos consists mainly of proteoid, restioid and ericoid fynbos with thicket in wetter areas, and is found on gentle to steep southern and northern slopes of the relatively low Tsitsikamma mountain range. Structurally, it can be described as medium dense, tall proteoid shrubland over a dense moderately tall, ericoid-leaved shrubland. It occurs on acidic lithosol soils derived from Ordovician sandstones of the Table Mountain Group, but plinthic catenas are prominent (Mucina & Rutherford 2006). Important species include the Proteaceae, Leucadendron conicum , L. eucalyptifolium , L. uliginosum subsp. glabratum and Protea mundii (tall shrubs), together with the Erica discolor variant speciosa and E. sparsa . Herbs include Commelina africana and Gazania krebsiana subsp. krebsiana , together with the Graminoids Restio triticeus and Tetraria capillacea . Endemic taxa include Aspalathus teres subsp. thodei , Erica trachysantha and E. zitzikammensis , with the conservation status of the type described as “vulnerable” (Mucina & Rutherford 2006). Eastern Coastal Shale Band Vegetation occurs on narrow, linear shale bands on clayey soils derived from the Cedarberg Formation. Important taxa include the widespread Leucadendron eucalyptifolium and Protea neriifolia , as well as L. salignum and Leucospermum cuneiforme . The type has been classified as “endangered” (Mucina & Rutherford 2006). A more detailed classification of the fynbos vegetation is provided by Vlok et al . (2008).

Natural forest in the Tsitsikamma has been classified as Southern Afrotemperate Forest (Mucina & Rutherford 2006), comprising of mountain forest, coastal scarp and plateau forest (Geldenhuys 1993). Mountain forest lies scattered on the southern mountain slopes in fire

management plan GRNP Appendices 39 refugia where they are protected against bergwind fires (Geldenhuys 1994). They are of the wet forest types, dominated by fire-adapted species such as Ocotea bullata , Cunonia capensis and Virgilia divaricata on the forest edge. The Plateau Forest, comprising mainly of wet, moist and medium-moist High Forest, are located on the southern boundary of the catchment where, in places, they for a natural buffer between the catchment fynbos and the commercial pine plantations.

Fauna has not been well described for the catchment. Typical fynbos species occur, but the diverse habitats should support a relatively rich fauna.

FYNBOS FIRE ECOLOGY

Fire is the most important disturbance agent in fynbos vegetation, and essential in maintaining biodiversity and natural ecological processes (Stock & Allsopp 1992, Van Wilgen et al . 1992), an important management objective with the Tsitsikamma Mountain catchment area. Of critical importance is fire frequency, fire season and fire intensity (Van Wilgen et al . 1992, Bekker 1994).

The minimum fire frequency is determined by the time for the vegetation to reach maturity and species to complete their life cycle (Van Wilgen 1981). Although this would vary between mesic and drier sites, fire frequency for the Tsitsikamma Mountains, as a general guide, has been set at 15 years (Southwood & De Lange 1984), but it could be shorter for southern slopes and longer for northern slopes (Seydack et al . 1986).

The time of the year at which fires occur naturally in fynbos is largely determined by climatic factors (Van Wilgen et al . 1992) and can have a marked effect on species response to fire, especially in terms of regeneration patterns and the subsequent floristic composition of mature fynbos (Van Wilgen & Viviers 1985, Le Maitre 1988, Van Wilgen et al . 1992). Summer and early autumn fires seem to be beneficial for most fynbos plant species in terms of post-fire regeneration, which is also consistent with natural fire ignition patterns (Van Wilgen et al . 1992, Bekker 1994). As reported by Van Wilgen (1987), though, fires in fynbos can occur in all months under suitable weather conditions. The months December to April, but preferably January to March, have been set for prescribed burning for the Region (Seydack et al . 1986). However, recent research findings indicate variable recruitment conditions throughout the year towards the eastern part of the Cape Floristic Region where climate is less seasonal (Heelemann et al . 2007). Winter and spring fires in the Tsitsikamma could thus be favourable for fynbos recruitment. Also, from the limited data available on fire occurrence in the Tsitsikamma, it seems that natural fires (i.e. lightning fires) are less seasonally constraint in comparison with western and inland mountains. (A research project is underway to gain better insight into the natural fire regime for the Tsitsikamma).

In addition to fire frequency and fire season, post-fire regeneration and plant species composition after fire could also be severely impacted on by fire intensity (Van Wilgen et al . 1992, Bekker 1994). For example, low intensity fires would benefit sprouting species, and species with shallow seed banks, while other could benefit from a high intensity fire. The fire

management plan GRNP Appendices 40 intensity would depend on the fuel load, the compactness and arrangement of fuels, fuel moisture content and the rate at which they burn (Van Wilgen et al . 1992, Teie 2003), as influenced by climatic conditions. The late summer, early autumn period, regarded as the natural fire season, usually has weather associated with higher intensity fires (Bekker 1994). As natural fires have the greatest likelihood of occurring during fairly extreme weather conditions, these are also the more favourable conditions for prescribed burning to maintain biodiversity (Seydack et al . 1986), although constrained by practical management considerations.

The ideal fire regime for the maintenance of natural diversity in fynbos is one of exhibiting a natural range of variability in respect of fire frequency, fire intensity and also fire season, creating numerous transient niches (Seydack 1992, Bekker 1994).

FIRE MANAGEMENT OPTIONS

The management challenge was to develop and implement a fire management system that would incorporate the different aspects of a natural fynbos fire regime to ensure the conservation of biodiversity and the maintenance of natural ecological processes, but within the constraints of fire protection considerations. The most appropriate fire management system should thus make appropriate provision for meeting objectives of biodiversity conservation, the implementation of appropriate fire protection measures, meeting requirements of the National Veld and Forest Fire Act, but must be cost-effective and implementable.

Fire management options considered for the Tsitsikamma Mountain are Scheduled Block Burning, Natural Fire Zone Management and Adaptive Interference Fire Management ( vide Seydack 1992). The traditional block burn system entails the division of the whole of the area into smaller units (blocks) which are burnt in a predetermined year to create an approximately equal-sized veld age mosaic. The system, however, is intensive and often interfered with by wild fires. With the Natural Fire Zone Management System, fires of natural origin (e.g. lighting and rock falls) are allowed to burn themselves out without interference, while man- made fires are excluded as far as possible. Although the Tsitsikamma catchment is large enough for such a system, fires cannot easily be retained within the boundary of the area to prevent damage to adjacent property e.g. commercial plantations. The management system that was opted for was the Adaptive Interference Fire Management system”, as described by Seydack (1992).

ADAPTIVE INTERFERENCE FIRE MANAGEMENT SYSTEM AS IMPLEMENTED FOR THE TSITSIKAMMA

The Adaptive Interference Fire Management system is a viable management option for the Tsitsikamma Mountain, considering the extent of the catchment area and the presence of indigenous forest that can partly serve as a natural fire barrier between fynbos and assets that

management plan GRNP Appendices 41 need to be protected. The system provides for the use of both natural (particularly lightning) and artificial sources of ignition. Fires will be controlled in space in time as demanded by the appropriate fire regime for Tsitsikamma mountain fynbos, fire risk to private property, veld age configuration, etc., following an annual assessment of fire risk and intervention required for biodiversity conservation. Particular attention was given to requirements and measures to protect commercial plantations against fire damage, without jeopardising the objectives of biodiversity conservation. The adaptive interference management approach would allow for a natural range of variability in respect of fire frequency, fire season and fire intensity in a cost- effective way, while also meeting responsibilities in terms of fire protection.

Considering the objectives of biodiversity conservation and fire protection, the extent and topography of the Tsitsikamma Catchment area, and the locality and distribution of assets such as commercial plantations in the landscape, two fire management zones where distinguished. This includes a Wildfire zone where lightning would be the primary source of ignition, and a Block-belt zone where fire protection would be the primary objective.

Wildfire zone

The Wildfire zone comprises the largest part of the Tsitsikamma Mountain (Figure 1). In this zone man-made fires are limited with lighting fires the main source of ignition. The extent of the area and the limited extent of transformation of the natural vegetation (apart from invader plants), should allow for a self-driven fire regime, based on natural climatic patterns in the area. As such, management intervention in this zone should be limited, and self-ignited fires should, as far as possible, be allowed to burn themselves out. This should ensure a fire regime for maintaining natural ecological processes and patterns, i.e. a fire regime characterised by natural spatio-temporal variability in terms of frequency, seasonality, intensity and size.

Although management intervention should be limited, fire management in the Wildfire zone is flexible. As required by an adaptive management approach an assessment would be conducted annually to decide on the most appropriate, pro-active measures in anticipation of wildfires. The spatial control of fires is thus pragmatically determined according to local circumstances of veld age configuration, accessibility, hazard to property, fire protection measures already in place, etc. This should lead to the delineation of three sub-zones within the Wildfire zone (vide Seydack 1992): • Areas where fires should not burn: These would mainly be immature veld, or where damage to property or run-away fires may be anticipated. Such areas would have priority for intervention during wildfires. • Areas where fires may be allowed to burn unhindered. This would mainly be inaccessible areas where no immediate threat to property exists. The anticipated rate and pattern of spread could be considered though and a pro-active action plan put in place, should the fire cross into another sub-zone. • Areas where fires should preferably burn within the foreseeable future. This would mainly include senescent veld that requires a burn for the revitalization of the fynbos, old veld that could pose a fire hazard, etc. Here unscheduled controlled burns could be considered.

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Thus, although fire management intervention should be limited in this zone, pro-active fire protection measures could be considered if required, and the fire regime may be supplemented with intentional fires when and where required. On the southern slope of the Tsitsikamma Mountain where the catchment borders commercial pine plantations, the need for block burns for fire protection and control is largely covered in the block-belt zone. However, additional block burns could be a consideration to the north where the catchment borders farmland, should the need be identified.

Block-belt zone

This zone is largely confined to the first two west-east running ridges in the catchment, just north of the commercial plantations, and only where indigenous forest does not form a natural buffer between fynbos and plantations. The objective with this zone is to effectively reduce the risk of damage to plantations by fires originating in the catchment area, through block burns for wildfire control, while retaining the biological integrity of the area as far as possible. The blocks were strategically placed following an assessing of natural fire distribution patterns in the Tsitsikamma.

The zone essentially consists of two parallel fire belts, subdivided into smaller blocks, burnt on a rotational basis (Figure 1, Table 1). The exact block boundaries will be fixed in the field as the system is implemented. Where the boundary between two blocks follows the mountain crest, the boundary could (where appropriate and practical) rotate between the southern and northern slope to lower the risk of spot fires. The block-burn system is further supported by strategically placed fire belts, as formally agreed between SANParks and MTO. In addition, controlled burns in commercial plantations bordering the catchment ( vide De Ronde 1982) to create a plantation buffer zone of low fuel load, could further reduce the risk of run-away fires and the ability to retain them.

Fire management prescriptions and schedule

The block burn schedule and spatial distribution of blocks would allow for the creation of a mosaic of different veld ages required for fire protection.

Fire frequency and spatial distribution of burns

All blocks east of the Groot River will be burnt on an eight year fire frequency to adequately provide for fire protection. The burning frequency for blocks west of the Groot River (Keurbooms and some blocks at Bloukrans) has been set at 12 years, i.e. closer to the natural fire frequency for the fynbos type but still adequate for fire protection. Adjacent blocks are to be burnt four years apart (six at Keurbooms and Bloukrans) to create a mosaic of different veld ages along a north – south and west – east gradient. The different blocks are to be burnt as clusters (Keurbooms, Bloukrans, Olifantsvlakte, Lottering/Elandsbos, Kleinbos, Plaatbos, Blouleliesbos, Witelsbos West, Witelsbos East, Woodlands) to protect specific plantation areas. The block burning schedule is provided in Table 2.

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Most of the fynbos in the Tsitsikamma catchment is currently (2009), less than 10 years of age, mostly as a result of two large fire that ravage through the area in 1999, 2005 and 2009. Older blocks to the east of Storms River (Witelsbos and Woodlands) and those at Keurbooms, have been scheduled first for burning. Also, most of the burning blocks of a cluster are even- aged, either young veld or older than 10 years. To create a veld age difference of four or six years (east and west of Groot River respectively) between adjacent burning blocks but not to delay the implementation of the system, certain blocks will initially be burnt at a younger veld age than prescribed, i.e. 4 or 6 years.

The above burning schedule would ensure a mosaic of veld ages in such a manner that, regarding each block pair, the veld age of one block would be four years or younger (six west of Groot River) at any given time.

Fire season

The months November to April were initially set for prescribed burning for the Tsitsikamma Mountains (Southwood & De Lange 1984), later revised to December to April, but preferably January to March (Seydack et al . 1986). However, recent research findings indicate variable recruitment conditions throughout the year towards the eastern part of the Cape Floristic Region where climate is less seasonal (Heelemann et al . 2007). Winter and spring fires in the Tsitsikamma could thus also be favourable for fynbos recruitment.

For the burning blocks in the block belt zone, season of burn has provisionally been set at November to April. However, considering the initial research findings of favourable recruitment following winter and spring fires, and the extent of prescribed burning to be conducted in some years, earlier burns could be considered to accommodate the full burning programme. To enhance biodiversity and heterogeneity, burning season for the same burning block should preferably rotate between early and late summer burns, rather than being seasonally fixed for successive burns. The dominant type of fynbos in a burning block, could also impact on the most appropriate time for the burn.

Burning conditions

As natural fires have the greatest likelihood of occurring during fairly extreme weather conditions, these are also the favoured conditions for prescribed burning to maintain biodiversity (Seydack et al . 1986). Thus, although the practical execution of a fire must be considered with the choice of a specific time to conduct a block burn, hot, dry periods should not be avoided at all costs. Also, rather than burn under cool, moist conditions for ease of fire control, more elaborate pre-burning precautions (e.g. wider tracer belts), should be considered to allow for a hotter burn.

Fire weather conditions for burning would thus have to be a compromise between those permitting control of the burn, but achieving a clean burn of the vegetation. Unburnt patches that could result from a too cool burn, should be burnt if it is considered to jeopardize the fire protection objective with the burn.

management plan GRNP Appendices 44

Adaptive management

The burning schedule formulated for block-belt zone needs to be adhered to as far as possible to ensure that the maximum veld age considered necessary for effective fire protection is not exceeded, and that the mosaic of divergent veld ages aimed at, is achieved. However, the adaptive management approach does require an annual assessment of the schedule for at least the forthcoming two or three years to e.g. provide for budgetary requirements, operational planning, block burn preparations, etc. Also adaptations to the schedule could be considered following a fire risk assessment and changes in the veld age mosaic and local circumstances.

MONITORING AND OTHER FIRE-RELATED MANAGEMENT INTERVENTIONS

Fire records and fynbos monitoring

The Adaptive Interference Fire management system requires that accurate records be kept of all fires that occur in the area. This would include fires in the wildfire zone as well as in the block-belt zone. Information to be recorded for each fire includes date of fire, cause of fire, size of fire, weather conditions at time of fire, etc. All fires should be mapped as accurately as possible, as veld age maps would form the basis for the annual assessment of fire control and protection measures required.

It is envisaged that a post fire fynbos monitoring programme be implemented in both the wildfire zone and the block-belt zone. This would largely entail monitoring of Proteaceae regeneration to assess the effectiveness or possible impact of the prescribed fire regime (in terms of rotation, fire season and fire intensity) on fynbos diversity and composition. Where required, known populations of rare and endangered plant species would also be monitored.

Although prescriptions in terms of the appropriate fire regime for the Tsitsikamma Mountain fynbos do exist ( vide Southwood & De Lange 1984, Seydack et al . 1986), these would be revisited through a research programme to better inform fire management requirements in terms of biodiversity objectives as well flexibility in terms of the implementation of fire protection measures.

Forest management

Forest is not a fire-prone vegetation type and it is only in exceptional circumstances and extreme weather conditions that fire penetrates the forest proper (Moll, 1983). Forest ecotones are of importance because of their protective and ecological function, biodiversity and the high ratio of ecotone communities to forest communities (Everard, 1992). Fire has an important ecological function in terms of the maintenance of these natural forest/fynbos ecotones, which, could be severely impacted on by an unnatural fynbos fire regime. A too short rotation, for example, will not allow for the recovery of damaged ecotones before the next fire, while a too long fire rotation can lead to a high fire intensity, which could destroy small forest patches.

management plan GRNP Appendices 45

To allow for natural disturbance of the forest edge and to maintain the fynbos/forest ecotone, the forest edge will not be artificially protected against these fires. However, protective measures could be necessary where the forest edge is heavily infested by invader plants, resulting in a too heavy fuel load and altering the structure of the forest edge. Monitoring could also be required of small forest patches located in the block-belt zone that would be exposed to a relatively short fire rotation.

Invader plant control

It is of critical importance that the invader plant control programme be aligned with the fire management system developed for the area. Regards the block-belt zone, this would entail alignment with the block burn schedule, in such a way that input required and success with control are optimised. The same would apply to the wildfire zone, although the fire regime for the area is not fixed and the invader plant control programme will have to be adapted accordingly.

Resource use

Some of the burning blocks in the Block-belt zone have been incorporated into the resource- use zone for the Garden Route National Park (Table 1). The controlled harvesting of fynbos products would thus be allowed in these areas, subject to harvest prescriptions to ensure sustainability and to minimise impact on the resource. The alignment of harvesting with the burning schedule would be required.

CONCLUDING REMARKS

The envisaged fire management system for the Tsitsikamma Mountain catchment, should ensure that objectives in terms of fire protection as well as biodiversity conservation are met, if correctly implemented in an integrated way with other fire protection measures in the Region. The Tsitsikamma Fire Protection Association should serve as the platform for this.

REFERENCES

Bekker, S.J. 1994. The management of fynbos mountain catchments. In: Van der Sijde, H.A. (Ed.). South African forestry handbook . Southern African Institute of Forestry. Pretoria. pp. 691-714. De Ronde, C. 1982. The control and use of forest fire. In: Jordaan, J.V. (Ed.). Saasveld 50 1932-1982 . Department of Environment Affairs, Pretoria. pp. 193-198. Everard, D.A., 1992. Forest ecotone dynamics, problems and research priorities: a literature review. Deliverable Report FOR-DEA No. 496. Division of Forest Science and Technology, CSIR, Pretoria. 23 pp.

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Geldenhuys, C.J. 1993b. Composition and dynamics of plant communities in the southern Cape forests. Deliverable Report No. FOR-DEA 612. Division of Forest Science and Technology, Council for Scientific and Industrial Research, Pretoria. Unpublished. Geldenhuys, C.J. 1994. Berg wind fires and the location of forest patches in the southern Cape landscape, South Africa. Journal of Biogeography 21: 435-448. Heelemann, S., Proches, S., Rebelo, A.G., Van Wilgen, B.W., Porembski, S. & Cowling, R.M. 2007. Post-fire recruitment of non-sprouting serotinous Proteaceae in the eastern fynbos vegetation. Paper presented at the 30 th Fynbos Forum meeting with programme theme How well are we doing? Threatened habitat conservation: Targets and Management , 1-3 August, Langebaan. Le Maitre, D.C. 1988. The effects of parent density and season of burn on the regeneration of Leucadendron laureolum (Proteaceae) in the Kogelberg. South African Journal of Botany 54(6): 581-584. Moll, E.J., 1983. The Southern Cape Forests. South African Forestry Journal 127: 31-34. Van Daalen, J.C. 1988. Guidelines for the re-establishment of forest in the Southern Cape. Pamphlet No. 398. Department of Environment Affairs, Pretoria. 11 pp. Mucina, L. & Rutherford, M.C. (Eds.). 2006. The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria. 807 pp. Seydack, A.H.W. 1992. Fire management options in fynbos mountain catchment areas. South African Forestry Journal 161: 53-58. Seydack, A.H.W., Southwood, A.J., Bekker, S.J. De Lange, C., Swart, J.A.J. & Voges, K. 1986. Regional policy memorandum for the management of mountain catchment areas in the Southern Cape and Tsitsikamma forest regions. Department of Environment Affairs, Knysna. 59 pp. Unpublished. Southwood, A.J. & De Lange, C. 1984. Policy memorandum. Tsitsikamma Catchment Area. Department of Water Affairs and Forestry, Knysna. Stock, W.D. & Allsopp, N. 1992. Functional perspective of ecosystems. In: Cowling, R.M. (Ed.). The ecology of fynbos. Nutrients, fire and diversity . Oxford University Press, Cape Town. pp. 241-259. Teie, W.C. 2003. Fire manager’s handbook on veld and forest fires. Strategies, tactics and safety. South African edition. Southern African Institute of Forestry. Menlo Park. 488 pp. Van Wilgen, B.W. 1981. Some effects of fire frequency on fynbos plant community composition and structure at Jonkersberg, Stellenbosch. South African Forestry Journal 118: 42-55. Van Wilgen, B.W. 1987. Fire regime in the fynbos biome. In: Cowling, R.M., Le Maitre, D.C., McKenzie, B., Prys-Jones, R.P. & Van Wilgen, B.W. (Eds.). Disturbance and the dynamics of fynbos biome communities . South African National Scientific Programmes Report No. 135. Council for Scientific and Industrial Research, Pretoria. pp. 6-14.

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Van Wilgen, B.W. & Viviers, M. 1985. The effect of season of fire on serotinous Proteaceae in the Western Cape and the implications for fynbos management. South African Forestry Journal 33: 49-53. Van Wilgen, B.W., Bond, W.J. & Richardson, D.M. 1992. Ecosystem management. In: Cowling, R.M. (Ed.). The ecology of fynbos. Nutrients, fire and diversity . Oxford University Press, Cape Town. pp. 345-371. Vlok, J.H.J., Euston-Brown, D.I.W. & Wolf, T. 2008. A vegetation map for the Garden Route Initiative. Unpublished 1:50 000 maps and report supported by CAPE FSP task team.

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Table 1. Burning blocks comprising the block-belt zone in the Tsitsikamma

Cluster Block Fynbos type 2 Area Burning Veld age Comments N (ha) rotatio (years) o n (2009) (years) 1 Keurbooms 1 491.7 12 11 Dry fynbos with longer natural fire frequency; Zoned for resource use 2 354.6 12 11 Dry fynbos with longer natural fire frequency; Zoned for resource use 3 657.3 12 11 Dry fynbos with longer natural fire frequency; Zoned for resource use 4 1178.2 12 11 Dry fynbos with longer natural fire frequency; Zoned for resource use 5 715.1 12 11 Dry fynbos with longer natural fire frequency; Zoned for resource use Bloukrans 1 155.6 12 4 Lowland fynbos of high conservation value 2 145.8 12 4 Lowland fynbos of high conservation value 3 126.9 12 4 Lowland fynbos of high conservation value 4 386.2 12 4 Lowland fynbos of high conservation value 5 281.5 8 4 6 101.2 8 4 Olifantsvlakte 1 318.3 8 4 Zoned for resource use 2 110.5 8 4 Zoned for resource use 3 279.5 8 4 Zoned for resource use 4 125.4 8 4 Zoned for resource use Lottering/Elandsbos 1 237.4 8 4 Zoned for resource use 2 215.4 8 4 Zoned for resource use 3 206.6 8 4 Zoned for resource use 4 140.5 8 4 Zoned for resource use Kleinbos 1 169.1 8 4 Zoned for resource use 2 156.6 8 4 Zoned for resource use 3 227.4 8 4 Zoned for resource use 4 225.7 8 4 Zoned for resource use Plaatbos 1 233.8 8 11 In Nature Reserve zone 2 151.2 8 10 In Nature Reserve zone Blouleliesbos 1 178.2 8 10 Zoned for resource use 2 109.2 8 10 Zoned for resource use 3 195.1 8 11 Zoned for resource use 4 103.4 8 11 Zoned for resource use Witelsbos West 1 367.2 8 10 Zoned for resource use 2 311.5 8 13 Zoned for resource use 3 336.6 8 10 Zoned for resource use 4 336.0 8 10 Zoned for resource use Witelsbos East 1 175.9 8 10 Zoned for resource use 2 87.2 8 10 Zoned for resource use 3 143.7 8 10 Zoned for resource use Woodlands 1 45.8 8 10 Zoned for resource use 2 114.4 8 10 Zoned for resource use 1 Number and location of blocks to be finalised after the 2009/10 controlled burns 2 To be assessed and populated as the implementation of the system progresses

management plan GRNP Appendices 1

APPENDIX I

MANAGEMENT PLAN REFERENCES

A & MOUTON, PLEFN. 1999. A review of the amphibians and reptiles of the Cape Floristic Kingdom as indicators of centres of biodiversity, sensitive habitats and sites of special interest. Internal Report Internal Report contributing to the C.A.P.E Project, February 1999. Acocks, J.P.H. 1988. Veld types of South Africa. Memoirs of the Botanical Survey of South Africa 57. 146 pp. Africa: Environment & Wildlife. 2(4): 40-47. African savanna Afrixalus (Anura: Hyperoliidae). Unpubl. M.Sc. thesis. Allanson, B.R. & Herbert, D.G. 2005. A newly discovered population of the critically endangered false limpet Siphonaria compressa Allanson, 1958 (Pulmonata: Siphonariidae), with observations on its reproductive biology. South African Journal of Science 101(1-2): 95- 97. Allanson, B.R. & Howard-Williams, C. 1984. A contribution to the physio-chemical limnology of Swartvlei. Archive fur Hydrobiologie 99(2): 133-159. Allanson, B.R. & Whitfield, A.K. 1983. The Limnology of the Touw River Floodplain. South African National Scientific Programs Report No. 79. CSIR, Pretoria. 35pp. Allanson, B.R. Nettleton, J. & De Villiers, C. 2000b. Benthic macrofauna richness and diversity in the Knysna Estuary: a 50 year comparison. Transactions of the Royal Society of South Africa 55(2): 177-185. Allanson, B.R., Maree, B. & Grange, N. 2000. An introduction to the chemistry of the water column of the Knysna Estuary with particular reference to nutrients and suspended solids. Transactions of the Royal Society of South Africa 55(2): 141-162. Amphibians. South African National Scientific Programmes Report 151: iiv,1-242. An expedition to the Cape Province, October to December, 1940. Annals of the Transvaal Museum 20: 351-377. Angel, A., Brabch, G.M., Wanless, R.M. & Siebert, T. 2006. Causes of rarity and range restricyion of an endangered, endemic limpet, Siphonaria compressa . Journal of Experimental Marine Biology and Ecology 330(1): 245-260. Anon. 1985. Vegetation survey of the Tsitsikamma Coastal National Park. Forestry Branch, Department of Environment Affairs, Knysna. Anon. 2000. Guidelines for indigenous vegetation restoration following invasion by alien plants. Prepared for Working for Water Programme. Division of Water, Environment and Forestry Technology, Council for Scientific and Industrial Research, Pretoria. Anon. 2004. Botanical assessment of Erf 4016, Eastford, Knysna. Compiled for Withers Environmental Consultants. Nick Helme Botanical Surveys, Scarborough. Anon. 2005. The natural vegetation of the western Heads (Knysna). Notes on its ecological sensitivity and proposed future development. Regalis Environmental Services, Oudtshoorn. Arden-Clarke C. 1983. Population density and social organization of the Cape clawless otter (Aonyx capensis ) in the Tsitsikamma National Park. Msc thesis. University of Pretoria. Pretoria, R.S.A. 202 pp. Arden-Clarke, C. 1986. Population density, home range size and spatial organization of Cape clawless otter, Aonyx capensis, in a marine habitat. Journal of Zoology, London (A) 209: 201 - 211. Asaeda, T., Trung, V.K. & Manatunge, J. 2000. Modeling the effects of macrophyte growth and decomposition on the nutrient budget in shallow lakes. Aquatic Botany 68: 217-237 BAARD, EHW, BRANCH, WR, CHANNING, AC, DE VILLIERS, AL, LE ROUX, Barker, J. 1985. The effect of catchment land use on sediment input to Swartvlei. Unpublished MA dissertation. University of Cape Town. Barnes, K.N. (Ed). 2000. The Eskom Red Data Book of Birds of South Africa, Lesotho & Swaziland. BirdLife South Africa, Johannesburg, 168pp. BARNES, R. 2004. The distribution and habitat in the Knysna Estuary of the endemic South African mudsnail Hydrobia knysnaensis and the influence of intraspecific competition and ambient salinity on its abundance. African Journal of Aquatic Science 29(2): 205-211. BAUER, A.M., WHITING A.S. & SADLIER RA. 2003. A new species of Scelotes Bekker, S.J. 1994. The management of fynbos mountain catchments. In: Van der Sijde, H.A. (Ed.). South African forestry handbook . Southern African Institute of Forestry. Pretoria. pp. 691-714. Bekker, S.J. 1994. The management of fynbos mountain catchments. In: Van der Sijde, H.A. (Ed.). South African forestry handbook . Southern African Institute of Forestry. Pretoria. pp. 691-714. Bell, E.M., Lockyear, J.F., Mcpherson, J.M., Marsden, A.D. & Vincent, A.C.J. 2003. First field studies of an Endangered South African seahorse, Hippocampus capensis . Environmental Biology of Fishes 67: 35-46. Bennett, B.A., Attwood, C.G. & J.D. Mantel 1994. Teleost catches by three shore-angling clubs in the South-western Cape, with an assessment of the effect of restrictions applied in 1985. South African Journal of Marine Science 14: 11 - 18. Biodiversity: Western Cape Province, South Africa – reptiles. In: Western Cape Nature Conservation Board 2007. Biodiversity of the Western Cape 2007. Western Cape Nature Conservation Board, Cape Town. Biodiversity: Western Cape Province, South Africa – reptiles. In: Western Cape Nature Conservation Board 2007. Biodiversity of the Western Cape 2007. Western Cape Nature Conservation Board, Cape Town. Biology of the dominant fish species at Swartvlei. Fifth National Oceanographic Symposium, Rhodes University, Grahamstown. South African Journal of Science 79: 157pp. Biology of the dominant fish species at Swartvlei. Fifth National Oceanographic Symposium, Rhodes University, Grahamstown. South African Journal of Science 79: 157pp. BLAUSTEIN, A.R., ROMANSIC, J.M., KIESECKER, J.M.AND HATCH, A.C., Boltt, R.E. 1973. Coastal lakes benthos. Rhodes University, Institute for Freshwater Studies Annual Reports 1972/1973. Grahamstown.

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Butler, H.G. & R.F. Terblanche 1997. Investigation on the butterflies and moths (Lepidoptera) in the Tsitsikamma National Park. Research Report No. 1 of 1997. Progress Report for South African National Parks for 1996. 6 pp. BUTLER, J.R.A. AND DU TOIT, J.T. 1994. Diet and conservation status of Cape clawless otters in eastern Zimbabwe. South African Journal of Wildlife Research . 24 (3) 41 - 47. C.F.B., DE SA, R.O., CHANNING, A.,WILKINSON, M., DONNELLAN, S.C., RAXWORTHY, C., CAMPBELL, J.A., BLOTTO, B.L., MOLER, P., DREWES, R.C.,NUSSBAUM, R.A., LYNCH, J.D., GREEN, D.M., AND WHEELER, W.C. 2006. The amphibian tree of life.Bulletin of the American Museum of Natural History 297: 1-370. Calvo-Ugarteburu, G. 1998. The influence of environmental factors and disease on the biology and productivity of the Pacific Oyster Crassostrea gigas in the Knysna Estuary. Project Report, Department of Zoology and Entomology, Rhodes University, Grahamstown. 77pp plus appendices. Cameron M.J. 1982. Mountain and Forest Animals . Saasveld 50:1932-1982, 162-180. Directorate of Forestry. Department of Environmental Affairs. CAMERON, M. J., PRETORIUS, G. & REYNELL, D. G. (1978). Bestuursplan: Goudveld Staatsbos. Department of Environment Affairs, Forestry Branch, Knysna. (Unpublished). CAMERON, M.J. 1982. Mountain and Forest Animals. Saasveld 50:1932-1982, 162-180. Directorate of Forestry. Department of Environmental Affairs. Campbell, B.M. 1985. A classification of the mountain vegetation of the fynbos biome. Memoirs of the Botanical Survey of South Africa 50. 121 pp. CARRUTHERS, V.C. & ROBINSON, G.A. 1977. Notes on amphibia in the CARTER, B. (1970) Knysna Elephant Survey: February 1969–January 1970. Report to the Wild Life Protection and Conservation Society of South Africa, Eastern Province CARTER, N. 1971. The elephants of Knysna. Purnell, Cape Town. 220 pp. Cephalophus monticola populations in the Tsitsikamma Forest and Coastal National CHANNING, A. 1998. Tadpoles as bio-indicators of stream quality: A baseline study. CHANNING, A. 2001. Amphibians of central and southern Africa. Cornel University Chief Director Of Surveys And Mapping 1979. 1: 50 000. Map 3424AA Oubosrand. Private Bag, Mowbray. Chief Director Of Surveys And Mapping 1980. 1: 50 000 Map 3323 DC & 2423BB Nature’s Valley. Private Bag, Mowbray. Chief Director Of Surveys And Mapping 1981. 1: 50 000 Map 3323 DD & 2423BB Joubertina. Private Bag, Mowbray. Clarke BM, Hauck M, Harris JM, Salo K, Russell E 2002. Identification of Subsistence Fishers, Fishing Areas, Resource Use and Activities along the South African Coast. South African Journal of Marine Science 24: 425 – 437. CODY, M. L., BREYTENBACH, G.J., FOX, B., NEWSOME, A.E., QUINN, R.D. & W.R. SIEGFRIED 1983. Animal communities: diversity, density and dynamics. In Mineral nutrients in Mediterranean ecosystems (ed Day, J.A.) South African National Scientific Programmes Report 71, 91 – 110.

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Coetzee, D.J. & Pool, R.C. 1984. Stomach content analysis of the sea barbel, Galeichthys feliceps (Valenciennes in C & V), from the Swartvlei system, southern Cape. South African Journal of Zoology 20(1): 33-37 Coetzee, D.J. 1978. A contribution to the ecology of the zooplankton of the Wilderness Lakes. PhD. Thesis. University of Stellenbosch, Stellenbosch. 167pp. Coetzee, D.J. 1981a. Zooplankton distribution in relation to environmental conditions in the Swartvlei system, southern Cape. Journal of the Limnological Society of Southern Africa 7(1): 5-12. Coetzee, D.J. 1981b. Analysis of the gut contents of the needlefish, Hyporhamphus knysnaensis (Smith), from Rondevlei, southern Cape. South African Journal of Zoology 16: 14-20. Coetzee, D.J. 1982a. Stomach content analysis of the leervis, Lichia amia (L.), from the Swartvlei system, southern Cape. South African Journal of Zoology 17: 177-181. Coetzee, D.J. 1982b. Stomach content analysis if Gilchristella aestuaria and Herpestia breviceps from the Swartvlei system and Groenvlei, southern Cape. South African Journal of Zoology 17: 59-66. Coetzee, D.J. 1983. Zooplankton and environmental conditions in a southern Cape coastal lake system. Journal of the Limnological Society of Southern Africa 9(1): 1-11. Coetzee, J.C., ADAMS, J.B. & BATE, G.C. 1997. A botanical importance rating of selected Cape estuaries. Water SA 23: 81-93. Cotzee, D.J. & Palmer, N.G. 1982. Algemene fisiese en chemiese toestande in Eilandvlei , Langvlei en Rondevlei gedurende 1978. Bontebok 2: 9-12. CRAWFORD P.B., CRAWFORD S.A.H. & CRAWFORD R.J.M. (1983). Some CRAWFORD, R.J.M. & G.A. ROBINSON. 1984. History of the blue duiker CRAWFORD, R.J.M. 1982. Water mongoose Atilax paludinosus in the Tsitsikamma Coastal National Park. Koedoe 25: 121. CRAWFORD, R.J.M. 1984. Activity group structure and lambing of blue duikers Cephalophus monticola in the Tsitsikamma National Parks, South Africa. South African Journal of Wildlife Research. 14 (3) 65 - 68. Crewe, R.M. 1984. Report on the collection of insects and invertebrates at Harkerville . University of Witwatersrand, Unpublished Report. CSIR 1974. Proposed Braamekraal Marina, Knysna Model Studies. CSIR Report C/SEA 74/6 Stellenbosch. 29pp + 30 figs. CSIR 1976. Knysna Lagoon Model Investigation Part I – Main Report. CSIR Report C/SEA 7609. Stellenbosch. 43pp + 91 figs CSIR 1978. Hidrouliese studie van die Swartvlei Estuarium. CSIR Report C/SEA 7805/1. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch. CSIR 1981. Wilderness Report No. 1. Evaluation of prototype data and the application of a numerical model to the Wilderness lakes and Touws River floodplain. CSIR Report C/SEA 8113. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch.

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CSIR 1982. Wilderness Report No. 2. Evaluation of prototype flood conditions and application of the numerical model to conditions when the estuary mouth was opened . CSIR Report C/SEA 8255. Coastal Engineering and Hydraulics Division, National Research Institute for Oceanology, Council for Scientific and Industrial Research, Stellenbosch. DavieS, B.R. 1982. Studies on the zoobenthos of some southern Cape coastal lakes. Spatial and temporal changes in the benthos of Swartvlei, South Africa, in relation to changes in the submerged littoral macrophyte community. Journal of the Limnological Society of Southern Africa 8: 33-45. Davies, D.H. 1948. A new Goby from the Knysna River. Ann. and Mag. Nat. Hist. Series 12(1): 357-376. Day, J.H. 1967. The biology of the Knysna estuary, South Africa. In: Estuaries. Lauff, G.H. (Ed.). American Association for the Advancement of Science. Publication No. 83: 397-407. Day, J.H. 1981. Estuarine ecology with particular reference to southern Africa . Cape Town, Balkema. 411 pp. Day, J.H., MILLARD, N.A.H & HARRISON, A.D. 1952. The ecology of South African estuaries. Part III Knysna: A clear open estuary. Transactions of the Royal Society of South Africa. 33(3): 367-413. De Kock, A.C & Boshoff, A.F. 1987. PCBs and Chlorinated hydrocarbon insecticide residues in birds and fish from the Wilderness lakes system, South Africa. Marine Pollution Bulletin 18(7): 413-416. De Kock, A.C & Lord, D.A. 1986. Chlorinated hydrocarbon residues in African fish eagle (Haliaeetus vocifer ) eggs. Institute for Coastal Research Report No. 10. De Kock, A.C & Simmons, R. 1988. Chlorinated hydrocarbon residues in African marsh harrier eggs and concurrent reproductive trends. Ostrich 59(4): 180-181. DE Moor, F.C., De Moor, I.J., James, N.P.E. & Barber-James, H.M. 2004. An autumn survey of the freshwater macroinvertebrates in the Salt River, Southern Cape. Albany Museum Investigational Contract Report for Nature's Valley Trust. September 2004. DE VILLIERS, A.L. 1998. Threatened frogs. In: J.A. Harrison & M. Burger (Eds.). declines. Diversity and distributions 9: 123-140. Department of Environmental Affairs and Tourism. 2003. National Environmental Management: Protected Areas Act (Act 57 of 2003). Department of Environmental Affairs and Tourism, Pretoria. Department Of Water Affairs And Forestry (Dwaf) 1995. South African Water Quality Guidelines for Coastal Marine Waters. Volume 4: Mariculture. Pretoria. DIPPENAAR, M.J. 1994. Geographic Variation in Myosorex longicaudatus (Soricidae) Durrheim, G.P. 1993. Management Plan for Ysternek Nature Reserve. Conservation Planning Section, Chief Directorate: Forestry, Department of Water Affairs and Forestry Knysna. Durrheim, G.P. 2000. Timber yield regulation and harvest tree selection. In: Seydack, A.H.W., Vermeulen, W.J. & Vermeulen, C. (Eds.) Towards sustainable management based on scientific understanding of natural forests and woodlands. Proceedings: Natural forests and savanna woodlands symposium II, Knysna, South Africa, 5-9 September 1999. Department of Water Affairs and Forestry, Knysna. pp. 263-265.

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Durrheim, G.P. 2002. Plaatbos Nature Reserve Management Plan. Scientific Services Indigenous Forest Management Department of Water Affairs and Forestry. Duvenhage, I.R. 1983. Getyrivieroppervlaktes van sommige getyriviere aan die Kaapse kus . NRIO Internal Report, Stellenbosch. 172pp. Edge, D.A. & Pringle, E.L. 1996. Notes on the natural history of the Brenton Blue Orachrysops niobe (Trimen) (Lepidoptera: Lycaenidae). Metamorphosis 7: 109-120. edition. Struik. Cape Town. edition. Struik. Cape Town. ELDER, W.H. AND ELDER, N.L. 1970. Social groupings and primate associations of FAIRALL, N. & N. HANEKOM 1987. Population dynamics and possible management FAIRALL, N. & R.J.M. CRAWFORD 1983. Application of the Robson Whitlock Fijen, A.P.M. & Kapp, J.F. 1995. Wilderness Lakes catchment, Touw and Duiwe Rivers, water management strategy. Volume 1: Present situation. Department of Water Affairs and Forestry, Pretoria. 115pp Fijen, A.P.M. 1995a. Wilderness Lakes catchment, Touw and Duiwe Rivers, water management strategy. Volume 2: Water resources. Department of Water Affairs and Forestry, Pretoria. 57pp Fijen, A.P.M. 1995b. Swartvlei Lake catchment, Diep, Klein-Wolwe, Hoëkraal and Karatara Rivers, water management strategy. Volume 2: Water resources. Department of Water Affairs and Forestry, Pretoria. 47pp Filmalter E. & O’keeffe, J.H. 1997. Effects of land-use changes on the rivers on the Wilderness Lakes. Institute of Water Research, Rhodes University, Grahamstown. 96pp. FITZSIMONS, V.F.M., 1946. An account of the reptiles and amphibians collected on FRIEDMANN, Y. AND DALY, B. (EDS) 2004. Red Data Book of the mammals of Frogs and frog atlassing in southern Africa. First edition. ADU Guide 4: 64-66. from near Cape Town, Western Cape Province, South Africa. Proceedings of the California Academy of Sciences. 54(13): 231-237. from near Cape Town, Western Cape Province, South Africa. Proceedings of the California Academy of Sciences. 54(13): 231-237. FROST, D.R., GRANT, T., FAIVOVICH,. J., BAIN, R.H., HAAS, A., HADDAD, Geertsema, H. 1964. The Keurboom moth Leto venus Stoll Order: Lepidoptera (Hepialidae). Forestry in South Africa 5, 55–59. Geldenhuys, C.J. 1982. The management of the Southern Cape forests. South African Forestry Journal 121: 4-10. Geldenhuys, C.J. 1991. Distribution, size and ownership of forests in the southern Cape. South African Journal of Forestry 158: 51-66. Geldenhuys, C.J. 1991. Distribution, size and ownership of the southern Cape forests. South African Forestry Journal 158: 51-66. Geldenhuys, C.J. 1992. Disjunctions and distribution limits of forest species in the southern Cape. South African Journal of Forestry 161: 1-13.

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Hodgson, A.N., Allanson, B.R. & Cretchley, R. 2000. An estimation of the standing stock and population structure of Upogebia africana (Crustacea: Thalassinidae) in the Knysna Estuary. Transactions of the Royal Society of South Africa 55(2): 187-196. Holmes, P.M. & Richardson, D.M. 1999. Protocols for restoration based on recruitment dynamics, community structure and ecosystem function: Perspectives from South African fynbos. Restoration Ecology 7(3): 215-230. Howard-Williams, C. & Allanson, B.R. 1978. Swartvlei Project Report II. The limnology of Swartvlei with special reference to production and nutrient dynamics in the littoral zone. Institute for Freshwater Studies Special Report No. 78/3. Grahamstown. 280pp. Howard-Williams, C. & Allanson, B.R. 1979. The ecology of Swartvlei: Research for planning and future management. Water Research Commission, Pretoria. 26pp. Howard-Williams, C. & Allanson, B.R. 1981a. An integrated study on littoral and pelagic primary production in a southern African coastal lake. Archive fur Hydrobiologie 92: 507- 534. Howard-Williams, C. & Allanson, B.R. 1981b. Phosphorous cycling in a dense Potamogeton pectinatus L. bed. Oecologia 49: 56-66 Howard-Williams, C. & Davies, B.R. 1979. The rates of dry matter and nutrient loss from decomposing Potamogeton pectinatus in a brackish south-temperate coastal lake. Freshwater Biology 9: 13-21. Howard-Williams, C. & Liptrot, M.R.M. 1980. Submerged macrophyte communities in a brackish South African estuarine-lake system. Aquatic Botany 9: 101-116. Howard-Williams, C. 1977. The distribution of nutrients in Swartvlei, southern Cape coastal lake. Water SA 3: 213-217. Howard-Williams, C. 1978. Growth and production of aquatic macrophytes in a south temperate saline lake. Verh. Internat. Verein. Limnol. 20: 1153-1158 Howard-Williams, C. 1980. Aquatic macrophyte communities of the Wilderness lakes: community structure and associated environmental conditions. Journal of the Limnological Society of Southern Africa 6(2): 85-92. Howard-Williams, C. 1981. Studies on the ability of a Potamogeton pectinatus community to remove dissolved nitrogen and phosphorous compounds from lake water. Journal of Applied Ecology 18: 619-637 Howard-Williams, C., Davies, B.R. & Cross, R.H.M. 1978. The influence of periphyton on the surface structure of a Potamogeton pectinatus L. leaf (An hypothesis). Aquatic Botany 5(1): 87-91. Hughes, D.A. & Filmalter, E. 1993. Water quality management strategy for Wilderness, Swartvlei and Groenvlei Lake areas. Institute for Water Research, Rhodes University. Report to GFJ Inc. 57pp Hughes, D.A. & Görgens, A.H.M. 1981. Hydrological investigations in the southern Cape coastal region. Dept. of Geography, Rhodes University, Grahamstown. Hydrological Research Unit Report No. 1/181. 76 pp. Huizinga, P. 1985. Mathematical model study of the Knysna Estuary. The effect of river flow on salinity distributions. CSIR Report T/SEA 8506. CSIR, Stellenbosch. 13pp plus figures.

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Huizinga, P. 1987. Hydrodynamic model studies of the Swartvlei Estuary. CSIR Report T/SEA 8709. Stellenbosch. 22pp. in the Southern Cape Province, South Africa Journal of Mammalogy , Vol. 76, No. 4. pp. in the Tsitsikamma National Park, South Africa. Journal of Zoology, London 194: 467 – information on individual systems. Knysna (CMS13). Report No. 30. CSIR Research Report 429. 80 pp. information on individual systems. Knysna (CMS13). Report No. 30. CSIR Research Report 429. 80 pp. Jacot-Guillarmod, A. 1979. Report to the Lakes Area Development Board, George, on the invasive weed species in the area of the Wilderness Lakes. Institute for Freshwater Studies Special Report No. 79/3 : 20pp Jacot-Guillarmod, A. 1981. Structure of the floodplain vegetation with particular emphasis upon the status of exotic species. In: Jacot-Guillarmod, A. & Allanson, B.R (Eds.) Touw River Floodplain Report Part III. The chemical and biological environment and the impact of man. Institute for Freshwater Studies Confidential Report to the Council for Scientific and Industrial Research, Co-operative Scientific Programs. Jacot-Guillarmod, A. 1982. Checklist of the aquatic and floodplain vegetation of the Wilderness Lakes, southern Cape. Bontebok 2: 41-51. JANZEN, D.H. 1974. Tropical blackwater rivers, animals, and mast fruiting by the Dipterocarpaceae. Biotropica 6: 69–103. KIESECKER, J.M., BLAUSTEIN, A.R. AND BELDEN, R.K. 2001. Complex causes KINGDON, J., 1977. East African mammals III. Academic Press, London. KLOEPFER, D. (EDS) 2004. Atlas and Red Data book of the Frogs of South Africa, Lesotho and Swaziland. SI/MAB Series #9, Smithsonian Inst., Washington DC. Koen, J.H. & W. Breytenbach 1988. Ant species richness of fynbos and forest ecosystems in the Southern Cape. S.Afr. J.Zoo . 23 (3): 184 - 188. KOEN, J.H. 1983. Seed dispersal by the Knysna elephants. South African Forestry Journal 124:56–58. KOEN, J.H. 1984. A study of the distribution, population composition, movement and feeding of the Knysna elephants Loxodonta africana africana (Blumenbach 1797). Koen, J.H. 1985. Animal - Habitat Relationships in the Knysna Forest: Discrimination between Forest Types by Birds and Invertebrates. M.Sc. thesis, University of Cape Town. Unpublished Koen, J.H. 1988. Spatial distribution and resource partitioning of birds in the Knysna Forest, South Africa. African Journal of Ecology 26: 229-238 KOEN, J.H. 1991. The effect of rodent granivory on recruitment of the irregularly fruiting Podocarpus falcatus in the Southern Cape. South African Forestry Journal . KOEN, J.H., HALL-MARTIN, A.J., AND ERASMUS, T. 1988. Macro nutrients in plants available to the Knysna, Addo, and Kruger National Park elephants. South African Journal of Wildlife Research 18(2):69–71. Kok, H.M. & Whitfield, A.K. 1986. The influence of open and closed mouth phases on the marine fish fauna of the Swartvlei estuary. South African Journal of Zoology 21(4): 309-315.

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Whitfield, A.K. 1982. Trophic relationships and resource utilisation within the fish communities of the Mhlanga and Swartvlei estuarine systems. Ph.D. Thesis, University of Natal, Pietermaritzburg. 157pp. WHITFIELD, A.K. 1983. Effect of prolonged aquatic macrophyte senescence on the Whitfield, A.K. 1983. Effect of prolonged aquatic macrophyte senescence on the biology of the dominant fish species at Swartvlei. Fifth National Oceanographic Symposium, Rhodes University, Grahamstown. South African Journal of Science 79: 157pp. Whitfield, A.K. 1983. Effect of prolonged aquatic macrophyte senescence on the Whitfield, A.K. 1984. The effects of prolonged aquatic macrophyte senescence on the biology of the dominant fish species in a southern African coastal lake. Estuarine, Coastal and Shelf Science 18: 315-329. Whitfield, A.K. 1986. Fish community structure response to major habitat changes within the littoral zone of an estuarine coastal lake. Environmental Biology of Fishes 17(1): 41-51. Whitfield, A.K. 1988a. The fish community of the Swartvlei estuary and the influence of food availability on resource utilisation. Estuaries 11(3): 160-170. Whitfield, A.K. 1988b. The role of tides in redistributing macrodetrital aggregates within the Swartvlei Estuary. Estuaries 11(3): 152-159 Whitfield, A.K. 1989a. Ichthyoplankton interchange in the mouth region of a southern African estuary. Marine Ecology Progress Series 54: 25-33. Whitfield, A.K. 1989b. Fish larval composition, abundance and seasonality in a southern African estuarine lake. South African Journal of Zoology 24(3): 217-224. Whitfield, A.K. 1989c. Ichthyoplankton in a southern African surf zone: nursery area for the postlarvae of estuarine associated fish species? Estuarine, Coastal and Shelf Science 29: 533- 547. Whitfield, A.K. 1989d. Recruitment of ichthyoplankton into the Swartvlei Estuarine System. J.L.B. Smith Institute of Ichthyology Investigational Report No. 30: 5 pp. Whitfield, A.K. 1989e. The benthic invertebrate community of a southern Cape estuary: structure and possible food sources. Transactions of the Royal Society of South Africa 47(2): 159-179 Whitfield, A.K. 1992. Juvenile fish recruitment over an estuarine bar. Ichthos 36:23 Whitfield, A.K. 1993. Fish biomass estimates from the littoral zone of an estuarine coastal lake. Estuaries 16(2): 280-289 Whitfield, A.K., Allanson, B.R. & Heinecken, T.J.E. 1983. Estuaries of the Cape, Report No. 22: Swartvlei (CMS 11). CSIR, Stellenbosch. 62pp. Wildlife Research 13: 25 –26. Williams, M.C. 1996. Report on research findings concerning the life history and ecology of the Brenton Blue. Metamorphosis 7: 3-7. Wilson, V.J. & Child, G.F.T. 1964. Notes on bushbuck from a tsetste fly control area in northern Rhodesia. Puku 2: 118-128. Withers, I. 2001. Goodbye, big foot—or is it? Dramatic new evidence of more elephants in the Knysna forests. African Wildlife 55(1): 6–8. Witwatersrand, Johannesburg. 270 pp.

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