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Plant communities along the Eerste River, , : Community descriptions and implications for restoration

Authors: Riparian communities fulfil many functions, including the provision of corridors linking 1,2 Clifton S. Meek protected areas and other zones of high conservation value. These habitats across much of David M. Richardson2 Ladislav Mucina2,3 South Africa’s , especially in the lowlands, have been heavily impacted and degraded by human activities. There is increasing interest in the restoration of degraded Affiliations: riparian zones and the ecosystem services they provide to enhance the conservation value of 1Percy FitzPatrick Institute, landscapes. Previous studies of riparian vegetation in the Cape Floristic Region focused on DST/NRF Centre of Excellence, University of pristine headwater systems, and little is known about human-impacted communities that , South Africa make up most of the riparian vegetation in downstream areas. More information is needed on the composition of these plant communities to establish a baseline for management 2Centre for Invasion Biology, Department intervention. The riparian zone of the Eerste River in South Africa’s Western Cape province of Botany and Zoology, provides a good opportunity to study the features of riparian vegetation along the entire University, gradient, from pristine vegetation in a protected area through different levels of human- South Africa mediated degradation. Riparian vegetation was surveyed in 150 plots along the entire length 3Department of Environment of the Eerste River (ca. 40 km). Data were analysed using the vegetation classification and and Agriculture, Curtin analysis software package JUICE. Final groupings were plotted onto a two-dimensional University, Australia detrended correspondence analysis plane to check the position of the communities in the Correspondence to: reduced multidimensional space. Ten distinct plant communities were identified, including David Richardson several novel communities dominated by alien plant species. Descriptions of each plant community are presented. Diagnostic, constant and dominant species are listed and the major Email: structural and ecological characteristics of each community are described. [email protected] Conservation implications: Major changes to hydrological and soil properties, nutrient Postal address: Private Bag X1, Matieland dynamics and disturbance regimes and plant species composition along sections of the 7602, South Africa riparian zone mean that restoration of many of these habitats to their historic condition is not feasible. However, several native plant species that provide key ecosystem services persist in Dates: and adjacent to transformed communities, offering substantial opportunities for restoration Received: 20 June 2012 Accepted: 25 Oct. 2012 to achieve certain goals. Published: 15 Mar. 2013

How to cite this article: Meek, C.S., Richardson, D.M. Introduction & Mucina, L., 2013, ‘Plant The flora of South Africa’s Cape Floristic Region (CFR) is exceptionally diverse, with one of the communities along the Eerste River, Western Cape, highest levels of species diversity endemism of any flora in the world. Research on vegetation South Africa: Community ecology in the biome of the CFR has focused largely on aspects of the iconic Cape descriptions and implications shrublands – mainly fynbos and renosterveld (e.g. Cowling 1992; Cowling, Richardson & for restoration’, Koedoe 55(1), Art. #1099, 14 pages. Mustart 1997; Linder 2003). Riparian vegetation in the Fynbos biome is, however, structurally http://dx.doi.org/10.4102/ and compositionally distinct from the surrounding Cape shrublands. The predominant koedoe.v55i1.1099 vegetation type of riparian zones in the Western Cape has been variously named closed scrub Note: fynbos (Campbell 1986; Cowling & Holmes 1992; Cowling et al. 1997), hygrophilous mountain Additional supporting fynbos (Taylor 1978) and broad sclerophyllous closed scrub (Holmes et al. 2005; Kruger 1978). information may be found Other types, ranging from tall herblands to forests, may also occur in the riparian zone (Kruger in the online version of this article as an Online 1978). True afrotemperate forest may develop in areas of steep topography that afford protection Appendix: http://dx.doi. from fires (Manders & Richardson 1992). Despite being recognised as a special vegetation type org/10.4102/koedoe. in the biome, riparian vegetation has been paid very little attention in the regional literature. v55i1.1099-1. Where it is discussed, it is usually in the context of the larger terrestrial matrix. Very few formal classifications of riparian vegetation in the Fynbos biome have been published (but see Prins, Read online: Holmes & Richardson 2004; Sieben, Mucina & Boucher 2009; Sieben & Reinecke 2008) and none Scan this QR have looked at riparian communities outside relatively pristine headwater systems, which occur code with your smart phone or mostly in protected areas. mobile device to read online. Copyright: © 2013. The Authors. Licensee: AOSIS OpenJournals. This work is licensed under the Creative Commons Attribution License.

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Riparian communities in the CFR, especially those in the the town of Stellenbosch. At Stellenbosch, the river takes an south-western part of the region, have been heavily impacted abrupt turn, flowing southwards until it meets the Atlantic recently by human activities. Intensive land use in the Ocean in near the town of Macassar (Figure 1). The lowlands has led to increased soil erosion and sedimentation, river is influenced by flows from several tributaries along its increased nutrient inputs, and altered hydrological regimes route, with the bank-to-bank width averaging 5 m near the due to water impoundment and abstraction. In many cases, headwaters and 14 m in the lower river zone. The average vegetation clearance for agriculture and urbanisation has gradient of the river ranges from 24 m/km in the mountain included riparian vegetation. In addition, riverine corridors stream zone to as little as 2 m/km in the lower reaches of the provide ideal conditions for the establishment, proliferation river (King 1981). and spread of the many alien that were introduced to the region. The constant disturbance along rivers as a result The geology of the Eerste River catchment is composed of their dynamic hydrological nature and their ability to of Sandstone in the upper reaches and function as conduits for dispersal of propagules means that Malmesbury Shale and Cape Granite of the Bokkeveld riparian zones are severely invaded worldwide (Foxcroft, Group in the lower reaches. Low-lying coastal plain areas Rouget & Richardson 2007; Hood & Naiman 2000; Planty- are composed of aeolian sands (Brown & Dallas 1995). The Tabacchi et al. 1996). Positive feedback mechanisms promote catchment of the river lies entirely within a Mediterranean- rapid spread of aliens along riparian corridors through like climate zone, which has cool, wet winters and hot, dry human-induced habitat alteration and increased propagule summers. Annual rainfall averages range from more than pressure (Richardson et al. 2007). 3000 mm (Sieben et al. 2009) in the upper reaches of the river to around 570 mm in the lower reaches. About 80% of This combination of pressures on the natural environment has this precipitation falls in winter downpours, with only 7% led to the establishment of novel ecosystems (sensu Hobbs of the annual rain occurring between December and March et al. 2006) along the riparian corridors of the CFR. No (King 1981). corridors remain undisturbed in their lower reaches and it is unclear what the composition of historical plant communities As with most rivers in areas of concentrated development, in these downstream systems would have been (Brown 1998; the Eerste River has undergone dramatic changes in the Prins et al. 2004). These riparian ecosystems fulfil a range of years since European settlement. Intensive human use of the useful functions in the Western Cape and there is increasing river dates back to as early as 1697, with over-abstraction interest in conserving or restoring key ecosystem services in becoming a problem as early as 1862. Indigenous trees were these zones. Intact riparian corridors in the CFR have also been used for firewood and timber in the early days of European identified as being important for migration and exchange of settlement and were later replaced by plantings of English inland and coastal biotas in a human-dominated landscape in oaks (Quercus robur) (Brown & Magoba 2009). Currently, the the face of global change (Rouget et al. 2003). Before sensible landscape surrounding the Eerste River supports a number restoration efforts can be undertaken, more information is of land uses, including nature conservation, commercial required on the composition of these ecosystems. The Eerste forestry, residential use, various forms of agriculture River provides a superb natural laboratory for the study of (vineyards, orchards and crop production) and communal the changing dynamics of a river and its associated riparian grazing. habitats. Approximately the first 6 km of the upper river reaches This study aimed to describe the plant communities along are bordered by natural vegetation within the Hottentots the entire length of the riparian corridor of the Eerste River Holland and Jonkershoek Nature Reserves, and is relatively in the Western Cape province to establish a baseline for management and future research. The 150 permanent plots established in this study (see also Meek, Richardson & Areas of urban development ● Sampling points along the river Mucina 2010) form a permanent transect with considerable potential for other studies of biotic and abiotic changes along a well-studied river situated close to a university. Another Catchment objective of this work was to identify native species that could be used in restoration efforts in disturbed riparian areas in the Fynbos biome. Bottelary River Plankenburg Eerste River River Catchment Veldwagters River Stellenbosch Research method and design Eerste River Kleinplaas Study area Dam Kuils River

The Eerste River originates in the Jonkershoek Mountains in Eerste River N the Western Cape, South Africa. It has a catchment area of 420 km2, and it is approximately 40 km long. The source of Atlantic Ocean Macassar 5 0 5 10 Kilometres the river lies 60 km east of Cape Town at an altitude of 530 m, from where it flows in a north-westerly direction towards FIGURE 1: The location of the Eerste River in the Western Cape, South Africa.

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 3 of 14 Checklist unaffected by human influence. Although forestry plots. The decision on the plot size was based upon values plantations of Pinus radiata exist in portions of the landscape suggested in Westhoff and Van der Maarel (1973). in the Jonkershoek Nature Reserve, ‘a wide buffer zone of natural fynbos vegetation has been maintained between the All plant species within a plot were recorded, and height river and plantation areas’ (Meek et al. 2010). Since 1981, the and absolute cover for each species was estimated. Species upper reach of the river has been regulated by the Kleinplaas were determined to be native or alien following the criteria Dam, above which the Eerste River is perennial (Department of Pyšek et al. (2004) and using published floras including of Water Affairs and Forestry [DWAF] 2004). During summer Goldblatt and Manning (2000), Henderson (2001) and months, a municipal weir above the dam diverts the river’s Bromilow (2005). Species that could not be identified in the flow to the Ida’s Valley Dam, which supplies drinking water field were collected and labelled, and identifications were later made at the Bolus Herbarium, . and water for domestic use to the town of Stellenbosch (Brown All species determinations were verified by the herbarium & Dallas 1995). In order to provide compensation releases curator (Meek et al. 2010), and collections have been archived for downstream users, this flow is replaced with water from at the Centre for Invasion Biology at Stellenbosch University the through an interbasin transfer (http://academic.sun.ac.za/cib/). The nomenclature of the scheme. Water from the Theewaterskloof Dam contains high plant names follows Goldblatt and Manning (2000), except sediment loads, significantly reducing the water quality and for Searsia (see Moffett 2007). negatively affecting downstream biotas in the Eerste River (Brown & Magoba 2009). Surface soil samples collected from each plot were air dried and the pH was measured using a calibrated pH meter in 10-g ‘Stellenbosch (human population ca. 117,000) is the main samples of soil suspended in 25 mL of potassium chloride urban area along the river, with additional urban development solution. Soil cation concentrations of sodium, calcium, present in Macassar’ (Meek et al. 2010). Historically, the river magnesium, and potassium were measured using ammonium through Stellenbosch was notably shallower and more flood acetate extraction, and phosphorous was measured using the prone than it is today, containing heavily wooded banks and Bray II method for samples with a pH ≤ 6.9 (Bray & Kurtz numerous islands. Considerable canalisation of the river 1945) and the Olsen method (Olsen et al. 1954) for samples through the town of Stellenbosch has all but eliminated the with a pH > 7. Total carbon content (Walkley–Black method; river’s historic flood regime through this area (Brown & Walkley 1935) and electrical resistance were also measured. Magoba 2009). The banks of the river through Stellenbosch are currently both deep and heavily confined. Vegetation analysis and presentation Vegetation data comprising 150 sample plots were classified Just south of Stellenbosch, the Eerste River is joined by by beta-flexible clustering (β = –0.25), using square root the Plankenburg River. Water is abstracted at this point, transformation and Manhattan distances as the resemblance near the confluence with the Blouklip River, and (at lower measure to calculate distance between plots. The result levels by by owners of riparian properties) along most of the clustering was translated into a structured relevé of its length. Treated municipal effluent enters the river table using the software package JUICE version 7.0 (Tichý, through the Veldwagters tributary. Below the confluence Chytrý & Zelený 2009). Vegetation data recorded per vertical of the Veldwagters and Eerste Rivers, treated effluent is an layer were used to assure that both floristic and structural important component of river flow in summer. This makes components of vegetation composition would be considered the river largely perennial in downstream areas. by the classification. Fidelity values were calculated for each of the resulting clusters, and the species in each of the Field sampling columns of the synoptic table constructed by JUICE were sorted by fidelity using the phi coefficient of association Vegetation of the riparian zone was surveyed along the (cut level of 0.35; L. Tichý, pers. comm., 16 Mar. 2009). The entire length of the Eerste River, from the headwaters to just structured synoptic table was then translated back into a upstream of the estuary. All fieldwork was performed from relevé table and inspected for plausibility (interpretability) mid-September through mid-December (i.e. in spring) 2008 of the resulting clusters. Clusters were further split or in order to capture the greatest number of herbaceous species. combined until desired resemblance and homogeneity Vegetation plots were paired at intervals of 500 m along both criteria were reached. Interpretability of clusters with respect banks of the river and plot edges were extended to bankfull to field knowledge of the plant communities factored heavily margins. A total of 150 plots were surveyed and locations in the decision on the final make-up of clusters. The final 10 were recorded using a Garmin GPS Map76 handheld global clusters were plotted on a two-dimensional ordination plot positioning system. Plot sizes were adjusted depending upon of the detrended correspondence analysis (DCA; Hill & structure of the plant community at each plot location in order Gauch 1980) using the ecological software package Canoco to capture all regularly occurring species. Herb-dominated 4.5 (Ter Braak & Šmilauer 2002) to visualise the level of communities were sampled with plots of 20 m2, shrublands/ ‘spatial’ separation (relative to the ordination plane) of the thicket with 50 m2 plots, and forest communities with 100 m2 classification.

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Descriptions of each plant community were compiled. These stratum. This combination of tall tree and shrub strata creates list diagnostic, constant and dominant species (alien species a fully closed canopy above the forest floor. A herb stratum are marked using an asterisk), and provide a narrative (< 1.5 m), with an average cover of 29% and dominated by describing the major structural and ecological characteristics ferns, is the typical understorey of this community. Common of the community. Diagnostic species were determined understorey species include B. capense, B. punctulatum, using the phi coefficient of association between species and T. barbara and S. lanceum. Structurally and floristically, this vegetation units as a measure of fidelity (Chytrý et al. 2002). community is similar to the R. melanophloeos – C. capensis Species were considered to be diagnostic if they had a phi community described by McDonald (1988). It should be coefficient of 0.35 or higher. Constant taxa are those that noted that this community was completely devoid of alien occurred in at least 35% of relevés for a given community, vegetation, with the exception of a single Acacia melanoxylon whilst dominant species are those that had a cover value sapling present in one relevé. This relevé is adjacent to of 45% or greater in at least 5% of the relevés. A species a bridge used by forestry vehicles, suggesting that the binomial system was used in naming the communities, with A. melanoxylon seed may have been introduced into the one dominant and one diagnostic species being selected for habitat by a passing vehicle. the names. stellatifolium–Metrosideros angustifolia Results community Plant communities Abbreviated name: Brabejum community (Figure 2b). Ten distinct plant communities were identified. The floristic Diagnostic species: M. angustifolia, B. stellatifolium, Virgilia and structural composition of the communities is represented oroboides, I. mitis, P. aquilinum, Freylinia lanceolata, E. caffra, in the structured relevé table (Table 1). A list of all species Halleria elliptica, Rubus pinnatus, Psoralea aphylla, Phylica encountered in the vegetation plots appears as Online pubescens, Pentameris thuarii, P. calyptrate. Appendix 1. Constant taxa: Searsia angustifolia, Olea europaea subsp. –Brachylaena neriifolia community africana, africana, B. neriifolia, Prionium serratum, Abbreviated name: Cunonia community (Figure 2a). Pittosporum undulatum, D. glabra.

Diagnostic species: Cunonia capensis, Brachylaena neriifolia, Dominant species: B. stellatifolium, F. lanceolata, P. serratum. , Schoenoxiphium lanceum, Todea barbara, Blechnum capense, Cassine schinoides, acuminata, Aristea capitata, The Brabejum community can be found at altitudes from Rapanea melanophloeos, Metrosideros angustifolia, Podalyria 228 m to 400 m, on sandy soils with low pH (average = 4.3), calyptrata, Blechnum punctulatum, Brabejum stellatifolium, Erica low resistivity and low nutrient content. This community caffra, Ischyrolepis subverticillata, Pteridium aquilinum, Pellaea tends to be interspersed with the Cunonia community where calomelanos, Elegia capensis, Cliffortia cuneata, Diospyros glabra, it occurs at altitudes between 228 m and 262 m, above which Asparagus scandens, Cassytha ciliolate. the Cunonia community dominates. The amount of exposed rock ranges from 2% to 90%, with an average of 35%. The Constant taxa: M. angustifolia, I. mitis. vegetation is composed primarily of an almost impenetrable layer of shrubs (up to 5 m), with an average cover of 96%. Dominant species: C. capensis, R. melanophloeos, P. aquilinum, This stratum is dominated by B. stellatifolium, which occurs B. stellatifolium. in every relevé. Interspersed with the B. stellatifolium in this layer are M. angustifolia, F. lanceolata, I. mitis, E. caffra and The Cunonia community occurs at altitudes of 262 m – 496 m, P. calyptrata, amongst others. P. serratum often grows along primarily where there has been no disturbance on lands the water’s edge. Above this stratum there is sometimes immediately adjacent to the riparian zone. Rock cover an emergent tree layer (up to14 m) with an average cover estimates range from 10% – 80%, with an average of 42%. Very little soil is present amidst the rock and tree roots. of 11%. The dominant species in this stratum is V. oroboides. Where soils are present, they are sand or gravel with a low pH A herb stratum dominated by P. aquilinum provides the (average = 4.5), low nutrient content, and high resistivity (i.e. understorey in this community, with an average cover of low electrical conductivity). This community is characterised 15%. Whilst this community is composed primarily of native by a tall tree stratum (6 m – 13 m) with an average cover species, a number of alien species have managed to invade, of 72%. This stratum is dominated by C. capensis, I. mitis, including P. undulatum and Acacia saligna. Structurally and M. angustifolia and B. stellatifolium. Less common tree species floristically, this community has many similarities to the in this stratum include C. schinoides, Platylophus trifoliatus, K. africana – B. stellatifolium community described by Prins Maytenus oleoides and R. melanophloeos. A tall shrub stratum et al. (2004), the H. elliptica – B. stellatifolium community (3 m – 6 m) grows below this, and has an average cover of 61%. described by McDonald (1988) and the Restio gaudichaudianus It is dominated by B. neriifolia and B. stellatifolium, although – M. angustifolia community described by Van Wilgen and M. acuminata and M. angustifolia are also common in this Kruger (1985).

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 5 of 14 Checklist ...... + + + 2 3 2 3 3 1 2 ...... r + + + 1 2 4 2 2 2 1 ...... 1 2 2 5 6 ...... 2 2 5 7 1 ...... + + + 2 5 2 2 2 1 ...... r 2 4 2 5 6 3 ...... 3 4 2 2 5 2 3 ...... r r 4 2 2 8 1 ...... 3 2 3 5 2 2 4 2 0 3 ...... r + + + 2 2 2 2 1 1 2 6 ...... 5 5 2 9 4 ...... 5 5 1 5 ...... + 4 5 2 2 2 5 ...... r + 5 2 3 3 3 2 1 4 4 ...... r + + 5 2 2 2 2 7 5 ...... + 2 3 2 2 3 2 4 5 Table 1a continues on the next page → page on the next 1a continues Table ...... r r + + 2 2 1 2 7 7 ...... 3 5 1 1 ...... r 4 5 1 5 2 ...... + 5 3 1 ...... + + + + + 4 2 4 2 2 2 6 2 ...... r + + 5 1 2 1 6 0 1 ...... r r r + 3 2 2 3 1 2 2 3 3 ...... r r r + 2 3 1 2 4 0 2 ...... 4 2 5 2 % ...... r + 3 1 3 4 3 2 2 5 3 ...... 2 2 3 2 2 2 2 2 9 7 ...... + + 3 3 5 2 2 3 2 2 2 8 3 – 100 ...... 3 5 2 1 2 2 7 2 % ...... r + 2 2 4 2 3 2 2 1 2 7 4 ...... r + + 2 3 2 4 2 2 9 ...... r + + + + 2 2 2 4 1 9 ; 5, 76 ...... r 2 2 4 2 2 4 4 2 2 2 9 3 % ...... 3 4 1 5 2 4 1 ...... r r + + + + 2 2 2 2 4 2 1 2 2 6 4 – 76 ...... r r r r r 4 5 2 2 3 2 2 2 0 6 % ...... 4 3 3 2 2 6 1 ...... + 5 2 2 3 2 2 2 1 3 4 ...... 3 5 4 3 2 4 ; 4, 51 ...... + + + + 3 2 3 2 4 4 2 2 2 9 5 % ...... r r + + 2 4 2 4 1 1 3 3 2 4 2 ...... r r r r + 3 3 2 5 2 1 6 – 50 ...... r r r r + + 2 3 2 3 4 2 2 0 5 % ...... 2 2 3 2 4 4 2 1 2 0 1 ...... r r r + + 2 2 2 4 2 2 2 4 6 ...... r + + + 4 2 2 3 4 2 3 3 5 ...... Relevé r + + + + 2 1 3 5 2 2 2 1 2 9 6 ...... r r r + + 2 2 3 4 2 2 8 4 ...... + 2 2 4 5 4 2 9 – 25%; 3, 26 % ...... r r + + + 2 3 2 1 4 2 1 5 1 0 7 ...... r + + + 2 1 3 2 5 5 8 ; 2, 4 ...... + + 5 4 8 5 % ...... 5 2 2 5 5 5 ...... + + + 2 5 2 2 1 2 8 ...... ; 1, 3 r + 2 3 1 2 3 5 8 8 % ...... r r + + 3 2 5 6 8 ...... r r + 1 2 4 3 7 8 ...... r r + + + 2 2 2 4 1 2 2 2 2 7 ; +, 2 ...... r 2 1 4 8 % ...... r r + + 2 1 1 5 0 9 ...... r r + + 2 2 1 2 2 2 2 4 3 3 8 ...... r + + 2 3 4 1 8 ...... r r r r r r r + 2 2 5 3 6 6 ...... r + + + + + + + 2 1 2 2 4 2 3 3 6 ...... + 4 2 1 3 2 2 2 2 1 3 9 ...... r + + + + 1 3 2 2 4 2 2 1 1 3 2 1 4 ...... r + 2 5 2 2 2 1 3 ...... + 5 2 2 2 2 0 4 ...... r r r + + + + + 2 4 2 2 3 7 ...... r 2 4 2 2 2 2 5 1 ...... r + + + 2 2 2 5 4 ...... r + + 2 2 2 2 4 2 2 2 ...... + 2 4 2 4 2 7 3 ...... + 2 2 2 2 9 8 ...... r r r r r r + 2 2 5 ...... r r r + + 2 9 2 ...... r r + + + 2 2 3 3 2 ...... + 1 4 2 1 2 4 2 3 1 7 ...... 2 2 1 2 2 2 2 3 0 8 ...... r r 4 2 2 4 2 2 8 7 ...... 4 2 2 1 2 6 7 ...... + 4 2 2 2 2 2 5 7 ...... 3 2 2 4 7 ...... 2 4 2 3 1 2 3 7 ...... 2 2 2 2 3 2 2 2 2 8 6 ...... r + + 3 3 4 2 5 1 2 7 6 ...... r r + + + + + 2 1 2 1 2 2 1 5 6 Understorey Tree Understorey Tree Understorey Understorey Understorey Understorey Understorey Understorey Tree Shrub Understorey Understorey Understorey Tree Understorey Shrub Understorey Understorey Understorey Understorey Understorey Understorey Understorey Understorey Shrub Understorey Understorey Understorey Understorey Understorey Understorey Understorey Stratum † † † † † † † † † † † † † † † † † † † † † † † † † Structured relevé table containing taxa grouped according to their fidelity to particular communities. to particular their fidelity to according grouped taxa containing table relevé Structured , Alien species. For visual purposes, the percentage cover values for these taxa have been translated into modified Braun-Blanquet values as follows: r, 1 follows: values as modified Braun-Blanquet into been translated have these taxa for values cover visual purposes, the percentage For shading. by grey highlighted community are species of each Diagnostic † Cirsium vulgare Olea europaea ssp. africana Brachiaria deflexa Quercus robur Tradescantia fluminensis Tradescantia Pennisetum clandestinum Pennisetum Commelina benghalensis Galium spurium Sonchus oleraceus Species common to multiple communities multiple to Species common Tropaeolum majus Tropaeolum Populus x canescens Populus Populus x canescens Populus Populus community Populus Cyperus textilis Cynanchum obtusifolium Zantedeschia aethiopica Zantedeschia Salix mucronata ssp. mucronata Salix mucronata Salix community Raphanus raphanistrum Lycium ferocissimum Lycium Malva arborea Sonchus asper Lolium perenne Tetragonia fruticosa Tetragonia Rapistrum rugosum Rapistrum Persicaria lapathifolia Persicaria Picris echioides Rumex crispus Rumex Phragmites australis Phragmites Phragmites community Phragmites Geranium molle Hordeum murinum Medicago polymorpha Medicago Fumaria muralis Fumaria Bromus diandrus Ehrharta longiflora Ehrharta Erodium moschatum Erodium Bromus community Bromus TABLE 1a: TABLE Species

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 6 of 14 Checklist ...... + + 3 2 2 2 1 2 ...... Sporobolus africanus–Stoebe plumosa community + + 2 2 2 1 ...... + 6 ...... + 2 7 1 Abbreviated name: Sporobolus community (Figure 2c)...... 1 3 2 2 1 ...... 3 6 3 ...... r + 2 3 ...... 4 8 1 Diagnostic species: Stoebe plumosa, R. pinnatus, Sporobolus ...... + 0 3 ...... 4 1 2 2 6 africanus, Juncus lomatophyllus, Paspalum urvillei*, Passerina ...... 9 4 ...... 1 5 corymbosa, Searsia tomentosa, Cliffortia strobilifera, A. saligna*, ...... 2 2 2 5 ...... r + 3 4 4 Juncus effuses...... 7 5 ...... 2 4 5 ...... r + 2 2 2 4 7 7 ...... 2 3 1 1 ...... 2 2 5 2 Constant taxa: S. angustifolia, O. europaea subsp. africana, ...... 2 4 3 1 ...... 2 2 6 2 D. glabra...... r + 1 2 6 0 1 ...... + 4 1 2 3 3 ...... + 2 2 0 2 ...... r r 2 Dominant species: S. africanus. % ...... r 2 2 5 3 ...... 1 3 1 3 9 7 ...... + 2 2 3 8 3 – 100 ...... 3 2 7 2

% The Sporobolus community occurs along the banks of the ...... 2 7 4 ...... r 3 2 2 9 Kleinplaas Dam at an altitude of 277 m. It is represented ...... + 2 2 2 1 9 ; 5, 76 ...... 9 3 % ...... here by only two relevés. This community is dominated by + 2 4 1 ...... r r r r 2 3 2 2 2 6 4 – 76 ...... r r + 2 2 0 6 S. africanus, which forms its own stratum, 1.0 m high and % ...... 2 6 1 ...... 3 3 4 with an average cover of 53%. S. angustifolia and C. strobilifera ...... 2 4 ; 4, 51 ...... 2 9 5 emerge from this stratum, reaching 2 m in height, whilst % ...... r r r r + + 4 2 ...... r 3 1 6 S. plumosa and R. pinnatus form a low stratum up to 0.5 m. – 50 ...... r 2 2 0 5 % ...... + 2 2 0 1 Less prominent species found within this community include ...... r + 2 4 6 ...... r 3 5 P. urvillei, J. lomatophyllus and D. glabra, amongst others. Whilst ...... Relevé + + 2 3 2 9 6 ...... r + 2 2 2 2 8 4 P. serratum grows in dense patches along some areas of the ...... 2 2 9 – 25%; 3, 26

% Kleinplaas Dam, it was not captured in either of these relevés...... r + 2 1 0 7 ...... + + 5 8 ; 2, 4 ...... + 2 8 5 % ...... 2 3 5 5 ...... 2 2 8 Quercus robur–Cinnamomum camphora community ...... ; 1, 3 r r 1 8 8 % ...... r r 6 8 ...... Abbreviated name: Quercus community (Figure 2d). r 7 8 ...... r r 2 7 ; +, 2 ...... 4 8 % ...... 0 9 Diagnostic species: Cinnamomum camphora*, Ligustrum ...... r + 3 8 ...... + + 2 1 8 ovalifolium*, Acacia longifolia*, Syzygium australe*, Acacia ...... r r r 6 6 ...... r + + 3 6 elata*, Q. robur*, P. undulatum*, Quercus palustris*, Agapanthus ...... r + + 2 1 3 9 africanus (native, but here not in original habitat)...... r 1 1 4 ...... 1 3 ...... + 0 4 ...... r r r r 7 Constant taxa: B. stellatifolium, Commelina benghalensis*, ...... r + 2 5 1 ...... 1 2 1 2 2 5 4 Acacia mearnsii*, Tradescantia fluminensis*, O. europaea subsp...... + 2 2 2 2 2 ...... 2 2 7 3 ......

r africana, S. angustifolia, K. africana, P. serratum, M. angustifolia, 2 2 9 8 ...... r + 1 2 5 ...... r 2 2 9 2 Bromus diandrus*, Briza maxima*, Brachiaria deflexa, Ageratina ...... r + 2 3 2 ...... r 1 7 adenophora*...... + 2 2 3 0 8 ...... + 8 7 ...... + 6 7 ...... r 2 5 7 Dominant species: Q. robur*, P. serratum, B. stellatifolium...... r 4 7 ...... 1 2 3 7 ...... + + 2 3 2 1 8 6 ...... r 7 6 The Quercus community is the most prominent riparian ...... r r 4 2 2 5 6 plant community of urban areas in and around Stellenbosch, occurring at altitudes between 81 m and 225 m. Soils are sandy and mildly acidic (average pH = 5.08), with low resistivity and moderate nutrient content. Exposed rock ranges from 0 to 80 , with an average of 18 . This is the Shrub Understorey Understorey Tree Understorey Understorey Understorey Understorey Understorey Shrub Understorey Understorey Understorey Shrub Tree Understorey Stratum % % %

‡ most diverse of the Eerste River plant communities, thanks in part to the extraordinary number of alien species that have been introduced into the area. However, this community also maintains high native-species diversity relative to the Structured relevé table containing taxa grouped according to their fidelity to particular communities. to particular their fidelity to according grouped taxa containing table relevé Structured

† other alien-dominated communities. Absolute cover of † † alien species averages 123%, whilst the absolute cover of † † † † † native species averages 73%. Dominating the tree stratum † † † † † † (5 m – 21 m) in this community is Q. robur. The alien species † C. camphora, Eucalyptus camaldulensis and A. mearnsii play a secondary role in the canopy, as do the native species , Alien species. page. previous from continues , Data For visual purposes, the percentage cover values for these taxa have been translated into modified Braun-Blanquet values as follows: r, 1 follows: values as modified Braun-Blanquet into been translated have these taxa for values cover visual purposes, the percentage For shading. by grey highlighted community are species of each Diagnostic † ‡ Acacia mearnsii Acacia Vinca major Vinca Echium plantagineum Echium Acacia mearnsii Acacia Stellaria media Stellaria Xanthium strumarium Xanthium Oxalis pes-caprae Oxalis Briza maxima Briza Lolium perenne Olea europaea ssp. africana Bromus catharticus Euphorbia peplus Euphorbia Lactuca serriola Lactuca Sesbania punicea Eucalyptus camaldulensis Eucalyptus Senecio halimifolius Species common to multiple communities multiple to Species common TABLE 1a (Continues...): 1a (Continues...): TABLE Species O. europaea subsp. africana, I. mitis and M. angustifolia. Average

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 7 of 14 Checklist ...... + 4 2 0 5 1 ...... 2 2 9 4 1 ...... 3 2 1 7 4 1 ...... + 5 4 1 ...... 2 1 4 1 ...... 2 2 2 4 1 ...... 2 1 6 4 1 ...... 3 8 4 1 ...... 2 2 4 4 1 ...... 3 2 2 6 3 1 ...... + 3 1 5 3 1 ...... + 2 1 7 2 1 Table 1b continues on the next page → page on the next 1b continues Table ...... 3 2 2 3 0 4 1 ...... 2 2 9 3 1 ...... r r + 2 1 1 3 3 1 ...... 3 2 1 2 3 2 2 2 2 5 2 1 ...... 4 2 2 2 2 4 2 1 ...... r + 5 2 2 2 2 1 2 2 3 2 1 % ...... + 2 1 2 3 2 3 2 2 2 1 ...... r r r 3 3 2 1 2 2 1 2 1 – 100 ...... r + 2 2 2 2 8 3 1 % ...... r r + + 2 2 2 1 3 1 ...... r 4 2 2 2 4 3 1 ; 5, 76 ...... 3 1 3 2 2 3 1 % ...... 2 1 3 3 4 1 ...... – 76 + + 2 2 2 2 3 8 2 1 % ...... 2 4 2 7 3 1 ...... 3 2 2 2 8 0 1 ; 4, 51 ...... + 2 2 2 2 2 3 6 2 1 % ...... + 2 3 2 2 1 3 9 1 1 – 50 ...... % r + 1 2 2 2 2 1 2 1 3 1 2 2 2 5 1 1 ...... + 2 2 1 2 2 5 0 1 ...... + + 2 2 2 4 0 1 ; 3, 26 Relevé % ...... 2 2 2 2 1 2 2 3 1 1 ...... + 2 2 2 2 2 1 2 4 3 2 1 1 – 25 ...... + + 2 1 2 1 2 2 2 2 4 1 2 6 1 1 % ...... r r r + 2 1 1 2 4 4 1 1 ...... ; 2, 4 + 2 1 2 4 2 2 0 0 1 % ...... + 2 2 2 2 2 2 2 0 1 ...... 2 3 2 0 1 1 ; 1, 3 ...... r + + 2 3 8 1 1 % ...... 2 2 2 2 1 1 1 1 1 ; +, 2 ...... + + 2 2 1 1 4 2 9 9 % ...... + 2 4 2 5 9 ...... + 2 2 1 1 2 1 3 2 3 2 2 8 9 ...... + 2 4 ...... 1 3 ...... 4 8 ...... + 2 2 2 2 1 3 1 1 7 0 1 ...... r + 1 4 5 9 0 1 ...... + 6 9 ...... 4 4 2 5 4 3 ...... 2 1 5 6 5 ...... + 2 2 5 8 2 ...... + 2 2 5 9 1 ...... + + 2 2 3 0 2 1 ...... 1 2 2 3 4 4 9 ...... + 2 2 2 2 3 2 4 7 1 1 ...... 2 2 1 3 1 4 3 0 1 ...... r + + 2 5 1 0 1 ...... 5 7 9 ...... + 1 3 2 1 2 2 2 0 3 1 ...... + + + 2 4 1 9 2 1 Shrub Understorey Shrub Shrub Understorey Shrub Shrub Shrub Tree Shrub Shrub Shrub Shrub Shrub Shrub Shrub Shrub Shrub Shrub Tree Understorey Understorey Shrub Shrub Shrub Understorey Understorey Understorey Understorey Understorey Shrub Shrub Shrub Shrub Understorey Understorey Understorey Shrub Shrub Stratum † † † † † † † † † † † † † † † † † † Structured relevé table containing taxa grouped according to their fidelity to particular communities. to particular their fidelity to according grouped taxa containing table relevé Structured , Alien species. For visual purposes, the percentage cover values for these taxa have been translated into modified Braun-Blanquet values as follows: r, 1 follows: values as modified Braun-Blanquet into been translated have these taxa for values cover visual purposes, the percentage For shading. by grey highlighted community are species of each Diagnostic † Brabejum stellatifolium Brabejum and Cunonia community Brabejum Pentameris thuarii Pentameris Psoralea aphylla Phylica pubescens Phylica Rubus pinnatus Halleria elliptica Freylinia lanceolata Freylinia Ilex mitis Ilex Virgilia oroboides Metrosideros angustifolia Metrosideros Brabejum community Brabejum Quercus palustris Agapanthus africanus Agapanthus Cinnamomum camphora Pittosporum undulatum Pittosporum Syzygium australe Syzygium Acacia elata Acacia Acacia longifolia Acacia Ligustrum ovalifolium Ligustrum Cinnamomum camphora Quercus community Quercus Quercus robur Arundo and Quercus community Arundo and Quercus Anredera cordifolia Tradescantia fluminensis Tradescantia Rubus fruticosus Arundo donax Arundo community Paraserianthes lophantha Paraserianthes Cynodon dactylon Ageratina adenophora Sesbania punicea Cliffortia odorata Cliffortia Cliffortia community Cliffortia Juncus effusus Acacia saligna Acacia Searsia tomentosa Cliffortia strobilifera Cliffortia Passerina corymbosa Passerina Paspalum urvillei Paspalum Juncus lomatophyllus Sporobolus africanus Stoebe plumosa Stoebe Rubus pinnatus Sporobolus community Sporobolus TABLE 1b: TABLE Species

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 8 of 14 Checklist ...... 1 2 2 3 2 1 1 1 3 2 1 0 5 1 ...... + 1 2 2 2 1 1 2 2 2 2 2 2 9 4 1 ...... r + + + + + 2 2 2 2 1 2 2 7 4 1 ...... + 2 2 2 2 2 2 2 2 4 5 4 1 ...... r + + 1 1 3 2 2 2 2 2 2 3 2 2 2 1 4 1 ...... r r + + 1 1 2 3 2 3 2 2 2 2 4 1 ...... r + 1 2 2 2 1 2 3 1 2 1 2 2 2 3 2 2 2 6 4 1 ...... r + + + 2 2 2 2 3 4 2 8 4 1 ...... r r r + + + 2 2 1 2 2 2 3 2 2 3 2 2 2 4 4 1 ...... r r r r r r + 2 2 2 3 2 2 2 3 2 2 2 6 3 1 ...... r r + 2 3 2 2 2 2 2 2 2 2 2 2 1 5 3 1 ...... r + + 2 2 2 2 2 2 2 2 2 1 2 2 7 2 1 ...... r + + 2 2 1 2 1 2 2 2 1 0 4 1 ...... + 1 2 2 2 2 2 2 9 3 1 ...... r r + 2 1 2 2 2 3 3 2 2 2 2 2 3 3 1 ...... + + 1 2 1 2 1 2 1 5 2 1 ...... + + 2 2 1 1 4 2 1 ...... + 2 2 1 3 2 1 % ...... + 1 1 2 2 2 2 1 ...... + 2 2 2 2 2 1 2 1 – 100 ...... + 2 1 2 2 2 2 2 2 8 3 1 % ...... + 2 3 1 2 2 2 1 3 1 ...... + + 2 2 2 2 2 4 3 1 ; 5, 76 ...... + + 1 2 1 1 2 2 2 2 3 1 % ...... r + 4 1 2 2 2 3 4 1 ...... – 76 + 2 2 2 1 2 2 8 2 1 % ...... 2 7 3 1 ...... 1 2 8 0 1 ; 4, 51 ...... + + 2 3 2 2 1 2 2 2 2 6 2 1 % ...... r r 4 2 1 2 1 2 9 1 1 – 50 ...... % r + + + + 1 2 2 2 1 2 3 2 2 2 1 5 1 1 ...... + + + 2 2 2 1 4 2 5 0 1 ...... r + 2 2 2 2 4 2 4 0 1 ; 3, 26 Relevé % ...... r + 1 2 4 1 2 2 2 3 2 3 1 1 ...... + + + 3 2 2 2 1 2 2 2 2 2 1 1 – 25 ...... r r + + + + + 3 2 1 2 1 1 1 1 1 6 1 1 % ...... r r + 3 2 2 2 2 3 1 2 2 4 1 1 ...... r ; 2, 4 + 2 2 1 2 1 2 1 0 0 1 % ...... + 2 1 2 1 2 0 1 ...... + 2 2 1 2 2 2 1 1 0 1 1 ; 1, 3 ...... + 2 3 2 2 1 8 1 1 % ...... 2 2 1 2 1 3 1 1 1 1 ; +, 2 ...... r + + 2 2 2 4 9 9 % ...... + 1 2 2 2 4 2 2 5 9 ...... r r r + 2 1 2 1 2 2 3 8 9 ...... 2 2 2 2 2 2 4 ...... + 3 2 2 2 2 2 2 3 ...... + 3 2 2 2 8 ...... + + + + + 2 2 7 0 1 ...... + + 4 2 2 3 9 0 1 ...... + 2 1 3 6 9 ...... 2 4 3 ...... 3 1 2 6 5 ...... + 2 2 8 2 ...... 2 2 2 2 1 2 9 1 ...... 3 1 2 2 2 2 0 2 1 ...... r + + 2 2 2 2 2 2 2 2 2 3 2 4 9 ...... + 2 2 3 7 1 1 ...... + 2 2 2 3 0 1 ...... 2 2 1 0 1 ...... + 2 1 7 9 ...... 3 0 3 1 ...... 1 1 9 2 1 Understorey Understorey Understorey Understorey Shrub Understorey Tree Understorey Shrub Shrub Tree Shrub Tree Understorey Understorey Understorey Shrub Understorey Understorey Shrub Tree Shrub Shrub Shrub Understorey Shrub Understorey Shrub Understorey Understorey Understorey Shrub Tree Understorey Tree Tree Shrub Understorey Shrub Shrub Understorey Shrub Understorey Understorey Tree Tree Shrub Shrub Understorey Stratum ‡ Structured relevé table containing taxa grouped according to their fidelity to particular communities. to particular their fidelity to according grouped taxa containing table relevé Structured † † † † † † † † † † † † † † † , Alien species. page. previous from continues , Data For visual purposes, the percentage cover values for these taxa have been translated into modified Braun-Blanquet values as follows: r, 1 follows: values as modified Braun-Blanquet into been translated have these taxa for values cover visual purposes, the percentage For shading. by grey highlighted community are species of each Diagnostic † ‡ Pennisetum clandestinum Pennisetum Commelina benghalensis Galium spurium Brachiaria deflexa Acacia mearnsii Acacia Vinca major Vinca Acacia mearnsii Acacia Briza maxima Briza Eucalyptus camaldulensis Eucalyptus Sesbania punicea Eucalyptus camaldulensis Eucalyptus Senecio halimifolius Olea europaea ssp. africana Vicia benghalensis Digitaria sanguinalisv Digitaria Ipomoea purpurea Quercus robur Bromus diandrus Tropaeolum majus Tropaeolum Olea europaea ssp. africana Kiggelaria africana Kiggelaria africana Searsia angustifolia Prionium serratum Species common to multiple communities multiple to Species common Cassytha ciliolata Cassytha Diospyros glabra Diospyros Asparagus scandens Cliffortia cuneata Cliffortia Elegia capensis Pellaea calomelanos Pellaea Ischyrolepis subverticillata Ischyrolepis Cunonia capensis Brabejum stellatifolium Blechnum punctulatum Cassine schinoides Metrosideros angustifolia Metrosideros Rapanea melanophloeos Aristea capitata Aristea Maytenus Cassine schinoides Blechnum capense Brachylaena neriifolia Brachylaena Todea barbara Todea Schoenoxiphium lanceum Schoenoxiphium Ilex mitis Ilex Cunonia capensis Cunonia community Podalyria calyptrata Podalyria Erica caffra Erica Pteridium aquilinum Pteridium Brabejum and Cunonia community Brabejum TABLE 1b (Continues...): 1b (Continues...): TABLE Species

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 9 of 14 Checklist cover in this stratum is 67%. Below the tree stratum is a all the Eerste River plant communities, whilst the average diverse shrub stratum (1.5 m – 5 m) dominated by the natives absolute cover of native species is only 27%. Dominating the P. serratum and B. stellatifolium, with an average cover of 68%. tree stratum (5 m – 22 m) in this community is P. x canescens, Other native species that commonly occur in this stratum although E. camaldulensis or Q. robur replace P. x canescens are O. europaea subsp. africana, S. angustifolia, and K. africana. as the dominant species in some relevés. Other prominent Alien species common to this stratum include C. camphora, tree species in this community include A. mearnsii and Salix L. ovalifolium, A. mearnsii, A. longifolia and P. undulatum. babylonica. The only native tree species that seem to have Dominating the understorey are alien T. fluminensis and maintained a foothold in some areas are O. europaea subsp. Pennisetum clandestinum. Other common understorey alien africana and, to a lesser extent, K. africana and Salix mucronata mucronata species include B. maxima, C. benghalensis and A. adenophora. subsp. . The tree canopy is almost never closed in this community (63% cover on average), allowing for enough This stratum is sparser than the shrub and tree strata, with an light to promote growth of a herb stratum (< 1 m) with an average cover of 49%. average cover of 97%. Dominating this understorey are T. fluminensisand T. majus, both of which form dense mats of Cliffortia odorata–Ageratina adenophora community vegetation along the forest floor, which makes it difficult for Abbreviated name: Cliffortia community (Figure 2e). competing plants to germinate. Other common understorey species include B. deflexa, C. benghalensis and P. clandestinum, Diagnostic species: Cliffortia odorata, Sesbania punicea*, as well as the native species G. spurium and Z. aethiopica. The A. adenophora*, Cynodon dactylon, Paraserianthes lophantha*. shrub stratum (1 m – 4 m) is generally sparse (average cover of 22%) and is dominated by young P. x canescens. Constant taxa: P. clandestinum*, C. benghalensis*, F. lanceolata, B. diandrus*. a Dominant species: C. odorata.

A dense growth of C. odorata (up to 1.5 m) is the defining component of this community, which occurs at altitudes from 114 m to 225 m on sandy soils with moderate resistivity, low nutrient content and an average pH of 5.2. Often interspersed with C. odorata are the alien species A. adenophora, C. benghalensis, P. clandestinum and in several relevés, small saplings of S. punicea. Emergent through this ground cover are shrubs (up to 4 m) such as F. lanceolata and P. lophantha, providing a cover ranging from 0% to 74%, with an average of 35%. This community generally lacks a tree stratum, although in two relevés it contained a relatively open canopy of alien trees, including E. camaldulensis, Q. robur and A. mearnsii. This community is possibly identical b c to the C. odorata shrublands described by Sieben (2003).

Populus x canescens–Tropaeolum majus community Abbreviated name: Populus community (Figure 3a).

Diagnostic species: Populus x canescens*, Tropaeolum majus*. d e Constant taxa: T. fluminensis*, Galium spurium, P. clandestinum*, B. deflexa, Zantedeschia aethiopica, Sonchus oleraceus*, Q. robur*.

Dominant species: T. fluminensis*, T. majus*, P. x canescens*, Q. robur*, O. europaea subsp. africana, E. camaldulensis*, C. benghalensis*. FIGURE 2: Plant communities along the upper reaches of the Eerste River (e.g. primarily upstream of the Plankenburg River confluence): (a) Cunonia Of all the communities described here, the Populus community showing tall canopy of Metrosideros angustifolia with Brachylaena community is by far the most widespread along the Eerste neriifolia and Prionium serratum in the undergrowth; (b) Brabejum community showing shrubby Brabejum stellatifolium and Metrosideros angustifolia with River, accounting for close to a third of all relevés. It is found Prionium serratum growing along the river bank; (c) Sporobolus community primarily adjacent to agricultural lands at altitudes between with Sporobolus africana in the foreground and Cliffortia strobilifera in the background next to the Kleinplaas Dam; (d) Quercus community. Brabejum 6 m and 162 m. Soils are sandy and mildly acidic (average stellatifolium mixes with Pittosporum undulatum and Eucalyptus camaldulensis pH = 5.48), with low resistivity and high nutrient content. This in the shrub layer with a canopy of Quercus robur and Ilex mitis and (e) Cliffortia community showing dense growth of Cliffortia odoratawith Freylinia lanceolata community has the highest absolute cover of alien species of above.

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Bromus diandrus–Erodium moschatum community within which very few, if any, native species are able to Abbreviated name: Bromus community (Figure 3b). survive. The thicket of A. donax emerges above a herbaceous stratum (up to 0.6 m) of other alien species, including Diagnostic species: Erodium moschatum*, Ehrharta longiflora, V. major, T. majus and T. fluminensis, as well as the weedy B. diandrus*, Fumaria muralis*, Medicago polymorpha*, Hordeum native species G. spurium. The tree K. africana was the only murinum*, Geranium molle*. other native species found growing within this community. The alien herbaceous vine A. cordifolia is often found twining Constant taxa: P. clandestinum*, G. spurium, T. majus*, up the A. donax stalks, whilst Q. robur creates a canopy up Z. aethiopica, Echium plantagineum*. to 13 m tall above the A. donax thicket. The tree stratum has a relatively low cover (average 26%) compared to the herb Dominant species: B. diandrus*, P. clandestinum*, T. majus*, (44%) and shrub strata (95%). Rapistrum rugosum*. Phragmites australis–Persicaria lapathifolia community The Bromus alien grassland community occurs at altitudes Abbreviated name: Phragmites community (Figure 3d). from 6 m to 114 m in heavily disturbed areas along the river, primarily downstream of Stellenbosch and adjacent Diagnostic species: Phragmites australis, Persicaria lapathifolia*, to agricultural or communal grazing lands. It generally does Rumex crispus*, Helminthotheca echioides*, R. rugosum*, not occur as a continuous unit, but rather in isolated patches Tetragonia fruticosa, Lolium perenne*, Sonchus asper*, Lycium interspersed amongst other communities. Soils are sandy, ferocissimum, Malva arborea*, Raphanus raphanistrum*, with low resistivity, neutral pH (average = 6.11), and high C. strobilifera. nutrient content. This plant community is dominated by non- native grasses and herbs and is unique amongst the riparian Constant taxa: Z. aethiopica, S. mucronata subsp. mucronata, communities represented here for its lack of a shrub or tree P. clandestinum*, G. spurium, F. muralis*, C. benghalensis*, stratum. Common species found within this community Xanthium strumarium*, T. majus*, S. oleraceus*, E. plantagineum*, include the annual grasses B. diandrus, E. longiflora and C. dactylon. H. murinum, as well as the perennial grass P. clandestinum. Herbs common to this community include the aliens Dominant species: P. australis, P. lapathifolia*. E. moschatum, T. majus and E. plantagineum, as well as the native species G. spurium and Z. aethiopica. Dwarf versions The Phragmites community occurs within an altitudinal range of many of these species can be found in heavily grazed between 5 m and 10 m, primarily adjacent to agricultural and areas along the river, where a majority of the species have grazing lands in the lower reaches of the river, as well as some untransformed lands close to the river estuary. Soils been pruned to within a few centimetres off the ground. are sandy with low resistivity, neutral pH (average = 6.80), P. clandestinum does particularly well in these areas owing to and high nutrient content. Similar to the Arundo community, its creeping, mat-forming nature. Very little bare ground is the most prominent feature of this community is the dense present within this community, which has an average cover thickets (3 m – 5 m tall) formed by the dominant species, of 96%. in this case P. australis. This community differs from the Arundo community, however, in that it contains a greater Arundo donax–Tradescantia fluminensis community cover percentage of native species and is found further Abbreviated name: Arundo community (Figure 3c). downstream. The tall herb P. lapathifolia (up to 1 m) is often interspersed with P. australis, particularly in relevés further Diagnostic species: Arundo donax*, T. fluminensis*, Rubus from the estuary. In some areas, portions of the P. australis fruticosus*, Q. robur*, Anredera cordifolia*. thicket have been grazed down and P. clandestinum has taken hold, along with other non-native herbaceous species such as Constant taxa: T. majus*, Vinca major*, G. spurium. F. muralis, C. benghalensis and X. strumarium. This community often abuts the Salix community (see later) and, as such, Dominant species: A. donax*, T. fluminensis*, Q. robur*. S. mucronata subsp. mucronata occasionally overhangs the P. australis, providing minimal tree cover. The Arundo community occurs within an altitudinal range of 15 m – 53 m, adjacent to agricultural lands downstream Salix mucronata subsp. mucronata–Cyperus textilis of Stellenbosch. Soils are sandy, with low resistivity, low community pH (average = 4.45) and unusually high phosphorus and Abbreviated name: Salix community (Figure 3e). potassium content. Species diversity is particularly low, with a mean of nine species per 50 m2 quadrat. As with Diagnostic species: S. mucronata subsp. mucronata, Z. aethiopica, the Bromus community, this community occurs primarily Cynanchum obtusifolium, Cyperus textilis. in isolated patches distributed amongst other downstream communities. The defining component of this community is Constant taxa: T. fluminensis*, R. crispus*, R. rugosum*, the dense stands of the alien species A. donax (up to 6 m), P. clandestinum*, C. benghalensis*.

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Dominant species: S. mucronata subsp. mucronata, C. textilis, Owing to the extensive changes in soil nutrients and species Cyperus dives, T. fluminensis*, T. majus*. composition that have occurred, complete restoration of these areas to their historic species assemblages (where As mentioned earlier, this community often abuts the this is known) is not feasible, and may even be detrimental Phragmites community and is found in similar areas along the (potentially leading to further degradation) in some cases. lower reaches of the river, at altitudes from 5 m to 15 m. Soils Steps can, however, be taken to remove some of the more are similarly sandy, with low resistivity, neutral pH (average aggressive alien species that are well established, whilst also = 6.83), and high nutrient content. Native species dominate re-introducing key native species into some communities in this community, with an average absolute cover of 95% (Galatowitsch & Richardson 2005). Whilst it is impossible as compared with an average of 65% for alien species. The to know with certainty which native species dominated in native S. mucronata subsp. mucronata forms a moderately tall these communities prior to human intervention, the native (6 m – 11 m) canopy above an otherwise heavily herbaceous species that persist provide some clues. For instance, the community. Prominent understorey herbs include the Populus community, which now occurs along 30% of the alien T. fluminensis and native Z. aethiopica, with native Eerste River, has a number of native species that maintain C. obtusifolium occasionally twining through the branches of a tenuous foothold in this novel community. These include the S. mucronata subsp. mucronata. In some areas, S. mucronata the native trees S. mucronata subsp. mucronata, O. europaea subsp. mucronata tends to grow in dense, shrub-like patches. subsp. africana and K. africana, all of which provide valuable Thick clumps of C. textilis often intermingle with shrubby ecosystem services to both humans and wildlife (Holmes S. mucronata subsp. mucronata, creating a tangle of et al. 2008; Richardson & Fraser 1995; Venter & Venter impenetrable vegetation. a Ordination The results of the DCA ordination are graphically illustrated in Figure 4. The plant communities show very clear ‘spatial’ separation (with reference to the reduced multivariate space) along axes 1, 2 and 3, suggesting that the floristic-based classification was robust. Discussion This study provides the first description of riparian plant communities along the entire length of a river system in the CFR, including the many novel communities that now occur in the human-impacted riparian areas in this region. A study of this nature poses a particular challenge owing to the continually changing nature of many of these plant b c communities. This is especially true in urban and agricultural stretches of the river, which continue to be transformed by the proliferation of already present alien plant species and the incursion of new alien species.

The distribution of plant communities along the Eerste River is not determined by any single factor, but by the interaction d e of several environmental factors. It is difficult to separate the effects of environmental factors in a system that has been, and continues to be, so heavily transformed by human intervention. No single environmental component can be treated in isolation, although it appears that land use has played an important role in shaping the composition of plant communities. Those riparian areas where the landscape FIGURE 3: Plant communities along the lower reaches of the Eerste River (e.g. primarily downstream of the Plankenburg River confluence): (a) Populus surrounding the river has been most heavily transformed community. Tall Populus x canescens towers above a herbaceous understorey by human factors are home to many of the novel plant of Tropaeolum majus and Vinca major, with Olea europea subsp. africana and Kiggelaria africana in the middle shrub layer; (b) Bromus community communities dominated by alien species that are described with Tropaeolum majus and Bromus diandrus in the foreground; (c) Arundo in this paper, including the Bromus, Populus, Arundo, and community showing dense thicket of Arundo donax with Tropaeolum majus in the understory and Anredera cordifolia climbing the Arundo stalks; (d) Quercus communities. The relationship between alien plant Phragmites community showing dense thicket of Phragmites australis, with species and surrounding land use was addressed in a Salix mucronata subsp. mucronata and Pennisetum clandestinum present on the left and (e) Salix community showing bushy patches of Salix mucronata separate study (Meek et al. 2010). subsp. mucronata.

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1996). It is likely that the dense understorey of T. majus and B. stellatifolium, M. angustifolia, P. serratum, and F. lanceolata. T. fluminensis within the Populus community has suppressed This, combined with the observation that these two the germination of native seedlings of these species, but communities exist both adjacent to and interspersed with they seem able to thrive below a canopy of P. x canescens. each other, suggests that the Quercus community may have Consequently, active planting will be necessary to establish historically looked very similar to the Brabejum community. denser stands of native vegetation, but little maintenance As such, land managers interested in restoring portions of other than managing the invasive species may be necessary the riparian habitat dominated by the Quercus community following these initial restoration activities. Once native can consider the composition of the Brabejum community species have grown large enough to provide the necessary to discover suitable species for restoration activities. The soil stabilisation, P. x canescens and other alien trees could be most appropriate inventory of native species for restoration removed in order to provide a greater opportunity for native along a particular stretch of river could further be confirmed species to thrive. by scanning upstream and downstream vegetation for key native species in these areas. There is an even greater opportunity for restoration along portions of the river dominated by the Quercus community, The Quercus community exists as far downstream as the as a diverse array of native plant species persist amidst confluence with the Plankenburg River, below which the alien species here. In fact, there is considerable overlap the Populus community becomes dominant. Interspersed between native species in the Brabejum community and with the Populus community is the Arundo community, those in the Quercus community, including species such as which is relatively rare along the river. The reed species A. donax is highly invasive and has major impacts in many Mediterranean-type ecosystems (Brooks et al. 2004; Coffman, 129 6 129 129 6 129 6 6 6 130 6 130 130 130 Arundo a Ambrose & Rundel 2010; Gaertner et al. 2009). Efforts should Bromus Populus be made to remove this species whilst it still occurs along Sporobolus only a small section of the Eerste River. Arundo donax tends to 129 6 129 6 129 Brabejum 6

6 130

6 130 23 130 23 23 130 Phragmites 75 23 crowd out all other species, creating monospecific stands and 75 75 Salix 65 65 65 73 7368 65 73 22 68 2273 22 providing few, if any, of the ecosystem services that other 68 68 76 22 37 Cliffortia 7668 76 37 22 7671 37 97 71 71 97 Quercus 71 97 97 80 80 63 80 93 alien species may provide. 66 63 6780 93 4093 Cunonia 66 63 6667 63 67 40 4093 101 23 66 63 786774 7 401545 1 101 101 90 23 78 774 1545 1 7 1545 1 21 101 90 75 23 788374 78 74 7 1545 1 21 7590 23 83 74 21 21 94 120 8375 83 79 120 94 120 75 83 79 94 94 120 79 79 94 65 79 65 73 69 103 68 65 73 22 69 103 103 68 65 73 22 69 6948 26 20 103 68 7673 22 37 26 694748 17 2091 137 68 76 22 37 48 1764 204748 26 2091 117 137 Interspersed with both the Populus and Salix communities 7671 37 47 64 914748 17 97202491 33 117 137 137 139 7671 37 64 41 64 33 9724 33 3 117 117 137 139 71 41 31 64 2459 9724 33106 3 117 104 125 138 139 139 71 81 31 3141 59 9724 3106 3 104 125 104 121123125 138 132 147 139145 81 3159 59 46106 106 3 96 121123 104 138123125 138 132 147145 149145 Axis 2 80 81 59 46 10696 105 121123132 147134 132 149145 80 93 46 46 96 105 123 131 134 132 141149147135 150 149145 66 63 6780 4093 46 105 105 134 131 134 141136140144135146148150 66 63 6780 40 93 910 1012 105 131 131 141136140144134135 150 141140144135146148150 127 is the Bromus community. As with the Arundo community, 66 63 67 401593 910 2 5091060 1012 8 131 128146148142 141136140144135146148150 127 90 66 63 786774 7 401545 1 72 60 509106036 1018 2 107 124128 142 128 142 127146148 127 90 78 74 7 1545 1 7221 50 366 506012 101 8 107 102107 124 128 142 127 90 83 2 AXIS 78 74 772 1545 1 7221 126 6150 3612 14 114 102107 124 108 90 83 2 AXIS 78 74 7 45 1 7221 6 611201232 3014 102114 102 108124 902 AXIS 21 6 94 14 1232 30 18 114 116 102113 108 AXIS 2 AXIS 120 83 2 AXIS 21 61 632 39 14 92 114 98113 115108 83 2 AXIS 79 9430166112044433239 1418 11411699 115108 83 79 3953941654186112092444332 30 18 92 116 9811311699 98113111 115 79 53 1654 4443 94165412044433942 18 92 99 11699 98113111 115 this community is in need of active restoration along the 79 53 1654 444342 35 13 11199 111 126 143 42 35 42103 13 112 111 126 143 69 70 1342103 35253813 112 126 143126 143 69 70 26 103 11253813 112109 11295 126 143 6948 17 26 20 70 2538 103 5211252749 109110 11295 694748 17 262091 112749 5211382710951 110 95109110 13795 100118 64 4748 17 26 2091 33 52 51 5211117275149 109100118110 13795 100 4164 4748 17 202491 33 5211751 137 100118 119 139 3141 64 24 33 3 57 117 104 137 125 138119 139 Eerste River. The Bromus community exists as a result of 81 31 41 64 59 24 3310657 3 117 104 121123119125 138119132 139145 81 3141 5959 46 24 106106 3 5796 104121123125 138119132 147 139149145 81 31 59 46 106 3 96 105 104 121123125 134138 132 147 149145 81 82 59 46 106 96 105 121123131 134 132 141147135 150 149145 82 46 55 96 105 131 134 141136140147144135146148150 149 82 910 2 1955 105 131 134 141136140136144135146148150 127 9501060 55 2 19558 131 128 142 141136140144135146148150 127 72 509106036 1986 2 19558 107 124 128 142 136140144146148 127 extensive clearing of woody vegetation by landowners and 72 6 50910603612 86 2 5819 8 34 102107 124 128 142 127 AXIS 2 AXIS 72 6 86 6150603612583286 14 58348 34114 102107 108124 128 142 AXIS 2 AXIS 72 6 61 361232863014 58 11434 102113107 108124 AXIS 2 AXIS 85 6 3930 18 114116 115 108124 AXIS 2 AXIS 85 6 168861 4443123239 301418 92 11411699 98102113 108 AXIS 2 AXIS 53 32 115 85 8587 53 16885461 4443323930141892 11499116 98111113 115108 87 88 85 53 881654 4443423930 18 9235 11699 98111113 115 87 53 881654 44434239 3592 132856 99 98111 126115 143 70 54 285642 35132856 112 111 126 143 intensive grazing along portions of the lower river. This has 70 42 352538281356 112 126 143 84 70 112538271349 109110 11295 126 143 84 84 70 52111125273849 109110 11295 100118 84 52112551382749 109110 95100118 5211275149 109110 95 100118 57 52 51 100118 119 5757 119 57 119 left the river in these areas extremely vulnerable to erosion 82 82 55 82 1955 82 1955 86 1955 86 5819 34 86 58 3434 and sedimentation, adding to the river’s already poor water 85 86 58 34

8587 -1 88 8587 -1 88 -1

-1 88 8587 -1 88 87 -1 88 2856 285656 84 -1 6 2856 quality in downstream areas. Where this community occurs 8484 84 00 0 8 8 8 AXISAxisAXIS 1 1 1 in communal grazing lands, the appropriate species for restoration can be found amongst the native species in the -1 -1 129 -1 -1 6 129 -1 129 6 129 6 6

5 130 6 130 130 Arundo 0 84 8 b Salix community, primarily S. mucronata subsp. mucronata 0 AXIS 1 Bromus 8 87 AXIS 1 Populus and C. textilis. In these cases, the riparian area will need 97 Sporobolus 129 to be fenced or otherwise protected from grazing until the 6 129 6 129 Brabejum 6 6 85 130 101 6 130 23 130 129 88 23 130 Phragmites 75 23 plants are of such a size that grazing will not disturb them. 75 Salix 65 8665 73 68 65 73 22 103 82 68 73 22 37 Cliffortia 68 76 22 37 117 This could probably be achieved over one or two growing 76 37 765871 97 Quercus 7155 97 72 80 70 80 93 66 63 6780 6445 4093 106 Cunonia seasons, given the rapid growth of these species. Once these 66 63 23 67 57 4093 101 23 66 63 78677422 16 7 401545 1 94 101 90 23 78 74 1553 7 154512 1 21 101 7590 23 83 78 74 37 7 441545 1 21 7590 23 83 41 1821 94 120 75 83 79 5149 94 120 75 31 60 79 52 94 120102 species have established themselves along the riverbank, 59 9 3679 120 65 7 17 14 13 103 65 73 6110 433969 103 68 65 73 22 471 6991 26 111 103 137 68 65 73 2222 37 6948 426948 17 3826 20 103 105 68 7673 22 37 50 32 4748 17 26 2091 98 137 68 76 22 7137 64 3048 17351126 20 96 117 121 137 7671 37 5441 6421 47 17 972491 33 117 125 137 150 127 139 they will provide not only numerous ecosystem services 71 31414664 279724 33 3 113 117 104 123 104 138125 147138 145 139 71 81 3141 59 92 9724 33106 124 104 121128123125 136140144135138146148132 149 139145 71 81 31 46 97 109 3 96 131104 121134123125132142 141138 132 147 149145 80 68 81 93 59 46 106 96 126 108105 104 121123125 134138 132 147 149145 Axis 3 80 20 3 96 105 123 131 134 132 147 150 149145 63 80 65 4093 79 46 25 96 115 105 143 131 134 139141136140144135 148150 149 66 63 6780 40 93 40 33910 2 105 131 134 136140144146148150 127 AXIS 3 AXIS 66 67 101 141135146 66 63 67 401593 2 5091060 1012 8 131 128 142 141136140144135146148150 127 but also additional forage for cattle being grazed in the area. 90 66 63 786774 787 401545 1 72 24 509106036 1012 112 107 124 128 142 146148 127 90 78 74 807 1545 1 7221 6 6 506012 101 8 102107 124 128 142 127 90 83 2 AXIS 78 74 677 1545 1 7221 6 6150 3612 14 107114 102107 108 AXIS 2 AXIS 72 32 95 108124 90 83 78 74 7 4573 1 7221 266 94 611201232 301418 11499 114116 119102113 124 AXIS 2 AXIS 120 8 83 2 AXIS 21 6 39 14 92 100114 98113 115108 83 2 AXIS 79 941661 444332 1418 11411699 108 83 7479 539416546112044433239301418 92 116 99 98113 115 81 79 5394165412044433942 18 92 11699 98113111 115 7975 53 1654 444342 35 13 99 111 126 143 In other areas of the river where this community exists, the 42103 13 112 111 126 143 83 69 70 42103 35253813 112 126 143 69 26 103 11253813 109 11295 126 143 6948 17 26 20 70 103 5211252749 109118110 11295 76 694748 17 262091 5211382751 109110 13795 100118 64 4748 17 26 2091 33 5211117275149 109110 13795 100 4164 4748 17 202491 33 34 5211751 137 100118 119 139 3141 64 24 33 3 57 117110 130104 137 125 138119 139 81 31 41 64 59 24 33106 3 117 104 121123125 138119132 139145 appropriate species for restoration will depend on adjacent 81 3141 5959 46 24 106106 3 5796 104121123125 138119132 147 139149145 81 31 59 46 106 3 96 105 104 121123125 134138 132 147 149145 81 82 59 46 106 96 105 121123131 134 132 141147135 150 149145 46 55 96 105 131 134 141136140147144135146148150 149 82 910 2 1955 105 131 134 141136140136144135146148150 127 9501060 2 19558 131 128 142 141136140144135146148150 127 72 63 509106036 86 2 19558 107 124 128 142 136140144146148 127 72 6 50910603612 86 2 5819 8 34 102107 124 128 142 127 communities. Where it exists interspersed with the Populus AXIS 2 AXIS 72 6 61506036123286 14 58 8 34114 102107 108124 128 142 AXIS 2 AXIS 72 6 61 361232863014 58 11434 102113107 108124 AXIS 2 AXIS 85 6 3930 18 114116 115 108124 AXIS 2 AXIS 85 6 168861 4443123239 301418 92 11411699 98102113 108 AXIS 2 AXIS 53 32 115 8587 53 16885461 4443323930141892 11499116 98111113 115108 90 66 85 53 881654 4443423930 18 9235 11699 98111113 115 87 53 881654 444319564239 3592 132856 99 98111 126115 143 70 54 42 35132856 112 111 126 143 70 42 352538281356 112 126 143 community, the appropriate species for restoration will be 84 70 112538271349 109110 11295 126 143 84 70 28 52111125273849 109110 11295 100118 84 52112551382749 109110 95100118 5211275149 109110 95 100118 57 52 51 100118 119 5757 119 57 119 those listed earlier for the Populus community; where it exists 82 82 55 82 1955 82 1955 86 581955 34 8686 5819 34 86 58 34 adjacent to the Quercus community, the appropriate species 85 86 58 34

8587 -1 88 8587 -1 88

-1 88 8587 -1 88 87 -1 88 2856 87 282856 -1 2856 84 56 for restoration can be found amongst the native species in the 84 84 -1 5 0 8 0 0 AXIS 1 8 8 Brabejum or Cliffortia communities. 0 AXISAxis 1 1 8 -1 -1 -1 -1 FIGURE-1 4: Detrended correspondence analysis ordination of the riparian No other study from South Africa has shown such distinct vegetation0 plots showing positionsAXIS of 1the 10 plant communities along8 axes (a) 1 and 2 and (b) 1 and 3. AXIS 1 differences amongst riparian plant communities as we

http://www.koedoe.co.za doi:10.4102/koedoe.v55i1.1099 Page 13 of 14 Checklist have revealed in the current study. In fact, other studies of Kritzinger-Klopper and Donald Iponga for assistance and riparian communities (e.g. Prins et al. 2004) tended to show companionship in the field, Terry Trinder-Smith for help with an overlapping continuum of plant communities rather than identifying plant specimens, Christy Momberg and Mathilda a series of unique communities with distinct differences. This van der Vyver for administrative support, Lubomír Tichý is largely because previous studies were confined to riparian for help with the JUICE software program, and the many communities of headwater systems, where most native landowners along the Eerste River, who kindly allowed us vegetation is intact and there is little human interference. access to their land. C.S.M. also thanks the staff and students Invasion by alien plant species and a range of human- at the Percy FitzPatrick Institute for their support during mediated habitat modifications in downstream areas of the the study. CapeNature kindly provided the plant collection Eerste River have radically changed vegetation structure and permits. composition (Meek et al. 2010). This has created an extensive array of novel plant communities, effectively disrupting the natural continuous vegetation patterns through increased Competing interests local dominance and associated disassembly processes. The authors declare that they have no financial or personal However, even the two primarily native plant communities relationship(s) that may have inappropriately influenced (those dominated by Brabejum and Cunonia), limited to the them in writing this article. upper reaches of the river, were quite distinct on the axes of the DCA. Whilst these communities share many floristic Authors’ contributions similarities, they are structurally quite different. Had the species structural data layers been combined into a single C.S.M. (University of Cape Town) and D.M.R. (Stellenbosch layer, these two communities would likely have overlapped University) were responsible for writing of the manuscript, to a much greater extent in the diagram, and would not have C.S.M. was responsible for vegetation sampling and data been as distinct in the initial cluster analyses. Of the 10 plant analysis and L.M. (Stellenbosch University) assisted with the communities, the Brabejum and Cunonia communities were ordination analysis and classification of plant communities. also the most tightly clustered in the ordination diagram, D.M.R. provided overall project guidance. whilst the compositional attributes of other plant communities were quite diffuse. This attests to the ever evolving nature References of the downstream plant communities, which continue to be Bray, R.H. & Kurtz, L.T., 1945, ‘Determination of total, organic, and available forms of changed and shaped by human activities in the surrounding phosphorus in soils’, Soil Science 59, 39–45. http://dx.doi.org/10.1097/00010694- landscape, as well as the major influence of humans on the 194501000-00006 hydrological regime of the river. Bromilow, C., 2005, Problem plants of South Africa, Briza Publications, Pretoria. Brooks, M.L., D’Antonio, C.M., Richardson, D.M., Grace, J.B., Keeley, J.E., DiTomaso, J.M. et al., 2004, ‘Effects of invasive alien plants on fire regimes’, BioScience 54, Conclusion 677–688. http://dx.doi.org/10.1641/0006-3568(2004)054[0677:EOIAPO]2.0.CO;2 Brown, C., 1998, ‘The ecological status of Western Cape rivers investigated: Shock An important priority for biodiversity conservation in South survey findings’, African Wildlife 52, 27–28. Africa is the protection and management of riparian habitats, Brown, C.A. & Dallas, H.F., 1995, Eerste River, Western Cape: Situation assessment of the riverine ecosystem. Final report, June 1995, Southern Waters Ecological which have become increasingly valuable owing to the Research and Consultancy, CSIR, Stellenbosch. multitude of ecosystem services they supply. Appropriate Brown, C.A. & Magoba, R., 2009, ‘Rivers and wetlands of Cape Town’, Report No. TT 376/08, Water Research Commission, Pretoria. management of these riparian areas depends largely upon Campbell, B.M., 1986, ‘Montane plant communities of the fynbos biome’, Vegetatio knowledge of riparian plant communities so that informed 66, 3–16. http://dx.doi.org/10.1007/BF00044078 decisions can be made regarding their management and Chytrý, M., Tichý, L., Holt, J. & Botta-Dukát, Z. 2002, ‘Determination of diagnostic species with statistical fidelity measures’,Journal of Vegetation Science 13, 79–90. restoration. This study details the extensive changes in plant http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x species composition that have occurred in many portions Coffman, G.C., Ambrose, R.F. & Rundel, P.W., 2010, ‘Wildfire promotes dominance of invasive giant reed (Arundo donax) in riparian ecosystems of Mediterranean-type of the Eerste River’s riparian zone, and helps to further an climate regions’, Biological Invasions 12, 2723–2734. http://dx.doi.org/10.1007/ understanding of how riparian plant communities in the s10530-009-9677-z Western Cape are changing as a result of anthropogenic Cowling, R.M. (ed.), 1992, The ecology of fynbos. Nutrients, fire and diversity, Oxford University Press, Cape Town. activities. Furthermore, it provides key data on those native Cowling, R.M. & Holmes, P.M., 1992, ‘Flora and vegetation’, in R.M. Cowling (ed.), species that continue to persist within and adjacent to alien- The ecology of fynbos: Nutrients, fire and diversity, pp. 23–61, Oxford University Press, Cape Town. dominated communities and offers insights with regard to Cowling, R.M., Richardson, D.M. & Mustart, P.J., 1997, ‘Fynbos’, in R.M. Cowling, options for restoration. Additional research is needed to D.M. Richardson & S.M. Pierce (eds.), Vegetation of southern Africa, pp. 99–130, determine long-term goals for the riparian vegetation along Cambridge University Press, Cambridge. Department of Water Affairs and Forestry (DWAF), 1993, ‘Western Cape System the Eerste River. All stakeholders need to be consulted to Analysis. Water Quality, Volume 1: General’, Report No. P G000/00/2891, Cape establish achievable goals. 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