A vegetation description and floristic analyses of the springs on the Kammanassie Mountain , Western Cape

The Kammanassie Mountain is a declared mountain catchment area and a Cape mountain zebra Equus zebra zebra population is preserved on the mountain. The high number of springs on the mountain not only provides water for the animal species but also contributes to overall ecosystem functioning. Long-term conservation of viable ecosystems requires a broader understanding of the ecological processes involved. It was therefore decided that a classification, description and mapping of the spring vegetation of the Kammanassie Mountain be undertaken. A TWINSPAN classification, refined by Braun-Blanquet procedures, revealed 11 major plant communities that could be related to geological origin. Habitat factors associated with differences in vegetation include topography, soil type and grazing. Descriptions of the plant communities include diagnostic species as well as prominent and less conspicuous species of the tree, shrub and herbaceous layers. The results also indicate a high species richness compared to similar regions and the difference between plant communities of wet and dry springs. This data is important for long-term monitoring of the spring ecosystems as well as for the compilation of management plans.


Introduction
The Kammanassie Nature Reserve was established in 1978 to conserve the Cape mountain zebra Equus zebra zebra population present in the area and the natural ecosystems as part of the mountain catchment area.The area has a high species richness compared to similar vegetation types (Cleaver et al. 2003;Cleaver 2004).A large number of springs is found scattered throughout the reserve contributing to the high species richness and different ecosystem functioning while also providing drinking water to various animals, especially the Cape mountain zebra (Cleaver 2004).
A total of 53 springs have been located on the Kammanassie Mountain.During 1998, it was noticed that certain springs were drying up.This was cause for concern for the survival of Cape mountain zebra on the reserve.Therefore, spring monitoring was initiated on the Kammanassie Mountain during 1999.Flora occurring at flowing springs were thought to be unique and in danger of being lost should further springs dry up on the mountain.It was decided to determine the floristic composition of springs to give an indication of what flora is currently found at the springs and what species could be lost should further flowing springs dry up on the reserve.This information is important for future management purposes.
Furthermore, the Department of Water Affairs and Forestry (DWAF) abstract approximately 0.65 x 106 m 3 /a of groundwater from five production boreholes on the In order to understand ecosystems and the management thereof (Bredenkamp 1982;Bezuidenhout 1993;Brown & Bezuidenhout 2000), it is important to classify, describe and map the different vegetation types found in the spring areas of the Kammanassie Mountain (Van Rooyen et al. 1981;Brown & Bezuidenhout 2000).This data will also be used in the long-term monitoring of the spring ecosystems as well as for compiling a management plan.

The Study Area
The Kammanassie Mountain complex is situated between the towns of Uniondale in the east and De Rust/Dysselsdorp in the northwest and west (Fig. 1).The mountain is an inselberg within the Little Karoo between the Swartberg and Outeniqua Mountains.The total area of the mountain range, managed by the Western Cape Nature Conservation Board (WCNCB), is 49 430 ha of which 21 532 ha are privately owned and a declared mountain catchment.The remaining 27 898 ha are state land, of which 17 661 ha have been declared State Forest.The Kammanassie Nature Reserve is situated between 33°33'S-33°37'S and 22°27'E-23°01'E and was proclaimed a protected area in 1978 as an important water catchment area (Cleaver 2002).
According to Rebelo (1996) and Lubke (1996) the Kammanassie Mountain falls into the fynbos and thicket biomes with small pockets of the forest biome present in the kloofs on the southern slopes of the Kammanassie Nature Reserve (Cleaver 2002).
The average monthly rainfall and minimum and maximum temperatures for the study area are indicated in Fig. 2. The reserve receives rain throughout the year, with an average annual rainfall of ± 450 mm.(Hälbich & Greef 1995).The Kammanassie Nature Reserve is an eroded remnant of the Kammanassie mega-anticline (Hälbich & Greef 1995).The Kammanassie Mountain Range comprises almost exclusively the resistant quartz arenites of the According to Kotze (2001), the Peninsula Formation is a highly competent succession of medium to coarse-grained, thickly bedded, grey sandstone.The Nardouw Formation weathers more brownish, and thin shale intercalations are more common, than in the Peninsula Formation.As a result the Nardouw Formation is more ductile (Kotze 2001).
Soil generally forms a thin (<1 m) veneer of silty sands as a result of the steep slopes of the Kammanassie Mountain and predominantly quartzitic rocks.Locally, clayey soil occurs in association with weathered shale horizons and, in particular, the Cedarberg Formation (Cleaver et al. 2003).

Methods
Relevés were compiled at 52 of the known springs on the Kammanassie Mountain.A first approximation of the main plant communities was derived by applying the two-way indicator species analysis (TWINSPAN) (Hill 1979) to the floristic data set.Further refinement of the classification was achieved by Braun-Blanquet procedures (Bredenkamp et al. 1989;Kooij et al. 1990;Bezuidenhout 1993;Eckhart 1993;Brown & Bredenkamp 1994).The results are presented in a phytosociological table produced by using MEGATAB (Hennekens 1996a).Springs showing similar plant communities were plotted on an ArcView (Ver.3.2) Geographical Information System (GIS) generated map, to indicate their localities.
Plant taxon names conform to those given by Goldblatt and Manning (2000).These names may therefore differ from the plant species list used by the TURBOVEG database (Hennekens 1996b), which is based on the PRECIS database of the National Botanical Institute (NBI) in Pretoria, as on date 1997.Red data status follows Hilton-Taylor (1996).

Classification
The results are presented in a phytosociological table (Table 1

Description of spring plant communities
In the descriptions of the different spring plant communities, all species groups refer to Table 1.The position of the springs (spring numbers) with similar plant communities is indicated in with same colour and symbol in Fig. 3.

Blechnum tabulare-Hippa frutescens fern community
The springs with vegetation belonging to this community are mainly found on southern slopes on the Kammanassie Mountain.
Altitude varies between 837-1394 m above sea level.The soil is loamy to clayey with rock cover varying between 5-40%.All the springs of this community are perennial and undisturbed.
Diagnostic species include the dwarf shrub Hippa frutescens and the ferns Blechnum tabulare and Blechnum inflexum (species group A).
This community can be sub-divided into the following three sub-communities: The shrubs Helichrysum petiolare (species group C), Psoralea verrucosa and small fern Hymenophyllum tunbridgense (species group F) are prominent.Other prominent species include the cosmopolitan weed Spergularia rubra and the moss Jamesoniella species.

Hippia frutescens-Helichrysum petiolare shrubland
The Hippia frutescens-Helichrysum petiolare sub-community is found at springs Kamm/w/35 and Kamm/w/36 (Mannetjiesberg) which were both flowing during the period of the survey.
These springs occur on gentle slopes (1-2°) between 1343-1394 m a.s.l.where the aspect is south-southwest.Soil texture is sandyloamy and with small-to-medium rocks, with a coverage of 30-40 % of the soil surface.
Shrub cover is 70 % and herbaceous cover is 30 %.  Seven families dominate the flora found at springs on the Kammanassie Mountain.The two largest families are the Asteraceae with 41 species, reflecting 14.5 % of the total flora and the Poaceae with 25 species (8.8 % of the total flora) (Fig. 4).These are followed by much smaller, yet significant families, represented by the Cyperaceae with 19 species (6.7 % of the total flora), Restionaceae with eight species (2.8 % of the total flora), Juncaceae with six species (2.1 % of the total flora), Rosaceae with six species (2.1 % of the total flora) and Scrophulariaceae also with six species (2.1 % of the total flora) (Fig. 4).
Two red data species were found, representing 1 % of the flora of springs on the Kammanassie Mountain.The endangered Erica sp nov (Ericaceae) was sampled at sample plot 18 (Kamm/w/19) and the rare Agathosma affinis (Rutaceae) in sample plot 24 (Kamm/w/25).Both these two springs dried up before the KKRWSS started abstraction.
A total of 11 alien species or cosmopolitan weeds were found at springs on the Kammanassie Mountain.

Geological origin of spring plant communities
The springs within the Hippia frutescens-Helichrysum shrubland (plant community 1.1) are all springs emanating at a perched water table of the Nardouw/Cedarberg Formation, while springs within the Hippia frutesens-Helichrysum petiolare shrubland (plant community 1.2) are all emanating at perched water tables on the Nardouw/Kouga Formation.Springs of the Hippia frutescens-Ehrharta species grassland (plant community 1.3) has its geological origin at shallow springs emanating at perched water tables on Cedarberg shale and Peninsula Formation.Thus all the springs within the Blechnum tabulare-Hippa frutescens fern (plant community 1) have their geological origin at shallow, perched springs.
The Eragrostis plana-Juncus lomatophyllus grassland (plant community 2) has a mixed geological origin, emanating from the Nardouw Formation and the Kouga/Baviaanskloof Formation that is in close contact with the Bokkeveld Group.All these springs originates predominantly from the Nardouw Formation with Cedarberg shale contact.
The nine springs found in the Berzelia intermedia-Psoralea verrucosa shrubland (plant community 3) are all shallow and perched, with six emanating from the Nardouw/ Cedarberg Formation, one from the Nardouw/Kouga Formation, and two from the Peninsula Formation.
A total of five of the seven springs of the Cliffortia ilicifolia-Stoebe plumose shrubland (plant community 4) has a predominantly Nardouw/Kouga Formation geological origin.Other origins include the Nardouw Formation with combined lithological and structural control.Therefore, the dominant geological origin is from the Nardouw Formation.
The Ehrharta ramosa-Aspalatus kougaensis shrubland (plant community 5) springs have their geological origin from the Nardouw Formation.Springs from the Conyza canadensis-Conyza scabrida shrubland

Discussion
A total of 244 plant species were found at 52 springs on the Kammanassie Mountain, representing 145 genera and 71 families.Of the 71 plant families, one gymnosperm, eight families of pteridophytes (11 %), 12 families of bryophytes (mosses)(17 %), seven families of monocotyledoneaes (10 %) and 43 families of dicotyledons (60 %) were recorded for the spring vegetation.The high presence of Asteraceae (14.5 % of the total flora) corresponds to Goldblatt & Manning (2000) and was similar to the Vermaaks, Marnewicks and Buffelsklip Valley floristic composition which had 16 % Asteraceae present (Cleaver 2004).Cyperaceae (6.7 % of the total flora) with 19 species and Restionaceae (2.8 % of the total flora) were the second and third most dominant families present at the springs on the Kammanassie Mountain.
The ratio of 6:1 dicotyledon to monocotyledon species observed for the spring flora compares well to that of the spring flora at Fernkloof Nature Reserve (four dicotyledons to one monocotyledon species) also situated in Mountain Fynbos vegetation (Mostert 2003).This ratio of spring flora for Kam-manassie Nature Reserve (6:1) is also higher than the ratio of 3:1 for the Cape Floristic region as given by Goldblatt & Manning (2000).However, the species richness for the Kammanassie Mountain springs is 47 species/100 m² compared to the much higher 126 species/100 m² for Fernkloof Nature Reserve (Mostert 2003).A possible explanation for this could be that the springs of the Fernkloof Nature Reserve are mainly situated on moist southern slopes (Mostert 2003) whereas a large number of springs on the Kammanassie Nature Reserve are located on drier northern slopes.This, together with the higher average annual rainfall of 674 mm for the Fernkloof Nature Reserve, compared to the average annual rainfall of 450 mm for the Kammanassie Mountain can explain the higher species richness found at Fernkloof Nature Reserve.
The Cliffortia ilicifolia-Stoebe plumosa shrubland represents springs that dried up before 1993.These springs are also perched and comprise drier shrubland vegetation, with grasses prominent.Both the rare Agathosma affinis (Rutaceae) and endangered Erica sp nov (Ericaeae) occur within this plant community, making it an important plant community to manage and conserve.The Erica sp nov, an endangered species, also occurs within the Ehrharta erecta-Rhus pallens community.
Flowing springs and the ones that have water have very different plant communities to those that are dry.A difference in plant species composition was also found between springs that have been drier longer than others.The species composition of flowing springs differ strongly with those of drier springs in that ferns, mosses and water plants dominate these communities.
Springs that have been dry for a number of years, have a shrub, grass and restio dominance while springs that have dried up recently (within the last 3 years) show a distinct difference, with no shrub layer present and the herbaceous layer being dominant.
There is a strong relationship between spring geological origin and plant communities

Conclusion
No similar vegetation descriptions have previously been carried out on the springs of the Kammanassie Mountains.This research therefore provides valuable data on these ecosystems.
The plant species identified from this study can now be included into the Kammanassie Nature Reserve Management Plan and will result in a more comprehensive plant species list for the reserve.The localities and plant communities within which previously unknown, rare, and vulnerable plant species occur were identified during this survey.
The wetness of a spring, period of time the spring had been dry and a spring's geological origin have influenced plant community composition of the springs on the Kammanassie Mountain.
If more springs dry up on the mountain, plant communities at these springs could change over time and species diversity could be expected to decrease as is the case with the dry and wet communities of this study.Important water dependent plants will be lost and 'wetland' plant communities will be transformed into dry shrub and grass dominated areas.The time it takes for these changes in plant species composition to form another plant community will, however, have to be determined through long-term monitoring of the different springs on the Kammanassie Nature Reserve.
These vegetation surveys and descriptions provide baseline information that allows similar surveys to be conducted in future.These monitoring data could then be compared with this study to determine if changes/shifts in plant communities have occurred.Since a change in habitat conditions will result in the loss of not only 'wetland' plant communities, but a host of other organisms such as invertebrates, frogs and fish species that depend on these plants for their existence, will also disappear.This will result in a reduction of flora and fauna biodiversity.The WCNCB should therefore continue to monitor rare and vulnerable species found during this survey to ensure their survival by applying the correct management strategies.Monitoring of spring vegetation should continue to determine changes in plant communities over time related to springs drying up.This could form part of a cleaver.qxd 2004/10/05 10:02 Page 19
Fig. 3: Location of the vegetation communities of springs on the Kammanassie Nature Reserve.

Table 1
Phytociological table of the vegetation of springs on the Kammanassie Mountaincleaver.qxd2004/10/05 10:02 Page 24 Rock cover is low and is less than 5 % in most cases while the soil texture is loamy-clay.
cleaver.qxd 2004/10/05 10:02 Page 27 Diagnostic species include the shrubs Helichrysum petiolare and Cliffortia burchelli (species group C).The shrub Hippia frutescens and the fern Blechnum tabulare (species group A) are prominantly present.The shrub Psoralea verrucosa (species group F) is also conspicuous in this subcommunity.1.3Hippiafrutescens-Ehrhartaspecies grassland This sub-community is found at the following springs: Kamm/w/05 (Buffelsdrif), Kamm/w/11 (Wildebeesvlakte), Kamm/w/33 (Elandsvlakte) and Kamm/w/46 (Upper Diepkloof).The spring Kamm/w/05 dried up in December 2001 and Kamm/w/33 dried up in 2000.At the time of the survey Kamm/w/11 had water but was not flowing and Kamm/w/46 was flowing.Diagnostic species include the shrubs Pelargonium cordifolium and Rhus tomentosa, the grass Ehrharta species , the restios Isolepsis verrucosula and Juncus capensis and aquatic plant Eleocharis limosa (species group D).The shrub Hippia frutescens, ferns Blechnum tabulare and Blechnum inflexum (species group A) are also prominent.Other prominent local species include the shrub Helichrysum cymosum (species group B), restio Juncus lomatophyllus and grasses Panicum ecklonii and Pennisetum macrourum (species group E) and Watsonia fourcadei (species group F). (Fig. 4) 2. Eragrostis plana-Juncus lomatophyllus grasslandFig.4.Dominant plant families reflecting the percentage of total flora of springs on the Kammanassie Mountain.cleaver.qxd2004/10/0510:02 Page 28aspect is northeast to north-northeast.Diagnostic species include the shrub Salvia namaensis, the forb Pelargonium radulifolium, the creepers Dipogon lignosus and Cissampelos capensis and the sedge Mariscus congestus (species group M).The grass Ehrharta ramosa (species group H) and tree Rhus pallens (species group K) occur locally while the sedge Mariscus thunbergii is local- Cleaver et al. 2003).Therefore, although this spring stopped flowing at surface the water table at this spring is at a sufficient level to sustain the wetter plant community found at this site.Springs Kamm/w/08 (1135 m a.s.l.) and 22 (1404 m a.s.l.) also originate at the Cedarberg Shale contact between the Peninsula and Nardouw Formations and dried up in 1999.These two perched springs have a drier plant community, completely reliant on rainfall, as the water table is not within reach of the plants at such high-altitude springs.It is interesting that the plant community at spring Kamm/w/09 has not changed sufficiently to fall within a drier type community.Should the water table at this spring decline it could result in a change of the present plant community and further monitoring is essential at this spring to better understand the relationship between the water table and plant community present.