A comparison of fynbos and non-fynbos coenoclines in the lower Gamtoos River Valley,southeastern Cape,South Africa

Patterns in the relative importance of structural attributes and growth forms along fynbos and non-fynbos coenoclines were studied to test the hypothesis that there would be less structural variation in the former because the overriding influence of low levels of soil nutrients would be manifest in a great deal of structural convergence in fynbos. The coenoclines were ranged along identical environmental gradients of increasing altitude, rainfall and soil moisture and decreasing climatic variability. Results showed that along the entire fynbos coenocline vegetation was structurally a small-leaved sclerophyllous shrubland with a graminoid understorey and, usually, a large-leaved (proteoid) shrub overstorey. Fynbos structure was interpreted largely as a response to low levels of soil nutrients. Non-fynbos vegetation ranged from mixed succulent-sclerophyllous and spiny large-leaved thicket at lower altitudes to tall mesic forest at the upper end of the gradient. Non-fynbos structure was explained in terms of variations in soil moisture and climate. An analysis of the biogeographical affinities of sample floras at sites along the coenoclines showed that fynbos vegetation was dominated by taxa endemic to the Cape phytochorion, although phytochorological mixing was pronounced at the lower altitude sites. The level of local endemism in the fynbos coenocline was relatively high; nearly all endemics were Cape fynbos taxa and their incidence increased with increasing altitude. These data indicate that fynbos vegetation has had a lengthy history in the southeastern Cape and that high altitude sites would have comprised a refuge for Cape taxa during unfavourable climatic periods. Non-fynbos vegetation ranged from dry subtropical Tongaland-Pondoland thicket with a strong Karoo-Namib component to temperate Afromontane forest. Levels of endemism were lower than the fynbos coenocline and decreased with increasing altitude. The high number of karroid endemics found in both coenoclines at low altitudes suggests that karroid vegetation would have been more widespread in the past, probably during the last glacial which was considerably drier than the present Holocene interglacial.     

Disturbance regimes as determinants of seed banks in coastal dune vegetation of the southeastern Cape

Soil-stored seed banks of grassland, fynbos and thicket, all growing on calcareous dunes and each subject to different disturbance regimes, were examined. Seed banks were determined from counts of germinants from 50 soil cores from each type. Aboveground estimates of plant species cover in 10 1-m² plots were used in determining vegetation/seed bank similarities. There was no evidence for seed bank densities to be markedly higher in the most frequently disturbed community (grassland -4273 seeds/m²) than the least disturbed community (thicket - 3417 seeds/m²). Highest similarity between seed bank and above-ground vegetation composition in terms of species and growth form/life-span classes was recorded for grassland (CC = 50%). Lowest similarity (CC = 13%) was found in the less frequently disturbed thicket where no seeds of climax trees were recorded in the seed bank. A fynbos community on a north-facing (warm, dry) slope had intermediate-sized seed banks (1683 seeds/m²) with intermediate vegetation/seed bank similarity (CC = 46%). However, on the south-facing slope, which has a large post-fire ephemeral herb component, seed banks were larger (4518 seeds/m²) but less similar to above-ground vegetation (CC = 39%o). Ordination (DCA) of vegetation data from the four communities was different from an ordination of their seed bank data. Fynbos shrub species were absent from seed banks of both grassland and thicket, even though secondary succession proceeds from grassland, through fynbos to thicket. Their seed banks appear less persistent than those of European heath or Californian chaparral shrubs.     

Insect species richness tracking plant species richness in a diverse flora: gall-insects in the Cape Floristic Region, South Africa

The Cape Floristic Region (CFR) is one of the most plant-species-rich regions in the world. It is also a warm temperate region and hypothetically should have high gall-insect species richness, making it interesting to investigate the relationship between the insects of the region and the rich flora. The relationship between gall-insect species richness (GSR) and plant richness was investigated for the Fynbos and for representatives of vegetation of the whole CFR. Samples (of up to 600 plants per transect for Fynbos) of woody shrubs were investigated for the presence of galls. The species richness of these insects was quantified, as well as plant species richness for each transect. GSR for Fynbos was compared to global figures for GSR. Fynbos harboured significantly more gall-insect species than other CFR vegetation types. GSR was positively correlated with CFR plant richness. GSR also closely tracked plant richness in Fynbos. GSR was not significantly influenced by other variables (elevation and aspect), suggesting that plant richness per se was an important factor in generating GSR. Fynbos GSR is comparable to other sclerophyllous regions of high GSR globally, corroborating that this vegetation type is conducive to gall-insect diversification. There is likely to be a high percentage of gall-insect endemism in the Fynbos, as might be expected from the high host fidelity of this insect group. Received: 22 September 1997 / Accepted: 16 February 1998     

Insects affect relationships between plant species richness and ecosystem processes

This study examined whether insects can alter relationships between plant species diversity and ecosystem function in grassland communities, by (i) altering biomass across a plant diversity gradient, (ii) altering relative abundances of plant species, or (iii) altering ecosystem function directly. We measured herbivore damage on seminatural grassland plots planted with 1, 2, 4, 8, or 12 plant species, and compared plant biomass in a subset of these plots with replicates in which insect levels were reduced. Plant biomass and herbivore damage increased with species richness. Reducing insect populations resulted in greater evenness of relative plant species abundances and revealed a strong positive relationship between plant species richness and above-ground biomass. Reducing insects also changed the relationship between plant species richness and decomposition. Plant species mixtures and their relative abundances partially explained plant biomass results, but not decomposition results. These results suggest that insects can alter relationships between plant diversity and ecosystem processes through all three mechanisms.     

Plant characteristics determine insect borer assemblages on Protea species in the Cape Fynbos, and importance for conservation management

The influence of regional climate, biotope and host-plant variables on the frequency of occurrence of insect borers associated with infructescences of Protea species in the species-rich flora of the Cape Fynbos was investigated. Large samples of infructescences (n=1000) were collected on a seasonal basis and borers identified and quantified. Data were analysed using DECORANA and CANOCO so as to correlate environmental variables with borer occurrence. Distinct differences in frequency of occurrence of the various insect taxa were recorded on the various plant species studied. These differences were primarily accounted for by physical host-plant characteristics (infructescence and seed set variables), and secondarily, by biotope variables and climatic factors. Fynbos plant characteristics therefore play a major role in determining insect abundance. Plant diversity in the Fynbos is maintained by burning. These management burns, however, should not be applied too frequently or over areas large enough to extirpate any borer species. As these borers are excellent umbrellas for many other insects, their conservation covers Fynbos insect diversity in general.     

Effects of alien pine plantations on small mammal community structure in a southern African biodiversity hotspot

Commercial plantations and alien tree invasions often have substantial negative impacts on local biodiversity. The effect of plantations on faunal communities in the fire‐adapted fynbos vegetation of the Cape Floristic Region biodiversity hotspot is not yet well quantified. We studied small mammal community structure in alien Pinus radiata plantations and adjacent fynbos regenerating after clear‐felling of plantations on the Cape Peninsula, South Africa. Small mammal sampling over 1,800 trap‐nights resulted in 480 captures of 345 individuals (excluding recaptures) representing six species. Significantly more species, individuals (12 X) and biomasses (29 X) of small mammals occurred on recovering fynbos sites compared to plantations. This was commensurate with a higher diversity of plant growth forms, vegetation densities and live vegetation biomass. Only one small mammal species, the pygmy mouse (Mus minutoides), was consistently trapped within plantations. Fynbos sites were dominated by three small mammal species that are ecological generalists and early successional pioneer species, rendering the recovering fynbos slightly depauperate in terms of species richness and evenness relative to other studies done in pristine fynbos. We make three recommendations for forestry that would facilitate the restoration of more diverse natural plant communities and progressively more diverse and dynamic small mammal assemblages in a key biodiversity hotspot.     

Biogeography and species richness of endophagous insects associated with Proteaceae in South Africa

The endophagous insects associated with Proteaceae of the Cape fynbos were compared to endophage assemblages from more northern non-Capensis Proteaceae. Insects were collected from Proteaceae in the Cape on a regular basis and additional records obtained from insect collections. Northern samples were collected more opportunistically or records were obtained from collections or through personal communication. The Cape fynbos genus Protea is utilized by many more insect taxa than the non-fynbos Protea species. The fynbos Proteaceae has very few species in common with the northern Proteaceae, yet each has many of their own distinct species. This suggests that the fynbos endophage insect fauna is distinct from that of the other regions. It appears that the high diversity of host plants in the fynbos has contributed to generating high, local endophagous insect diversity.     

Dissecting the plant-insect diversity relationship in the Cape

It has been argued that insect diversity in the Cape is disproportionately low, considering the unusually high plant diversity in this region. Recent studies have shown that this is not the case, but the precise mechanisms linking plant diversity and insect diversity in the Cape are still poorly understood. Here we use a dated genus-level phylogenetic tree of the Cape plants to assess how plant phylogenetic diversity compares with taxonomic diversity at various levels in predicting insect diversity. We find that plant phylogenetic diversity (PD) is a better predictor of insect species diversity that plant species diversity, but the number of plant genera is overall as good a predictor as PD, and much easier to calculate. The relationship is strongest between biomes, suggesting that the relationship between plant diversity and insect diversity is to a large extent indirect, both variables being driven by the same abiotic factors and possibly by common diversification, immigration and extinction histories. However, a direct relationship between plant diversity and insect diversity can be detected at fine scales, at least within certain biomes. Diversity accumulation curves also indicate that the way plant phylogenetic diversity and the number of plant genera increase over spatial scales is most similar to that for insect species; plant species show a greater increase at large spatial scales due to high numbers of local endemics.     

Effects of time since fire on birds in a plant diversity hotspot

Global changes are influencing fire regimes in many parts of the world. In the Fynbos plant diversity hotspot (Cape Floristic Region, South Africa), fire frequency has increased in protected areas where the mean fire interval went from 12–19 to 6–9 years between 1970 and 2000. Fire is one of the main drivers of plant diversity in the Cape Floristic Region. Too frequent fires threaten the persistence of slow-maturing plant species, and such insights have led to the adoption of fire management principles based on plant responses. The effects of fire on Fynbos fauna are much more poorly understood, and have not generally been considered in depth in Fynbos conservation policies, planning or management. We assessed the response of bird communities to long-term fire-induced vegetation changes using space-for-time substitution. We studied bird communities, vegetation structure and plant functional composition in 84 Fynbos plots burnt between two and 18 years before. Ten of the 14 bird species analysed showed a significant change in their abundance with time since fire. We observed a significant species turnover along the post-fire succession due to changes both in vegetation structure and plant functional composition, with a characteristic shift from non-Fynbos specialists and granivorous species to Fynbos specialists and nectarivorous species.If current trends of increasing fire frequency continue, Fynbos endemic birds such as nectarivores may become vulnerable. Conservation management should thus aim more carefully to maintain mosaics of Fynbos patches of different ages. Future research needs to estimate the proportion of vegetation of different ages and patch sizes needed to support dependent fauna, particularly endemics.     

Montane plant communities of Fynbos Biome

The vegetation of the mountains of the Fynbos Biome is classified and described, mostly on the basis of vegetation structure and higher taxa. Various gradients can be recognized. A gradient in soil conditions runs from coarse textured, nutrient-poor soils in non-grassy fynbos (Mountain Fynbos) through grassy fynbos and grassy non-fynbos (Eastern Fynbos and Grasslands & Grassy Shrubland) to finer textured and less nutrient-poor soils in the non-fynbos Karroid & Renoster Shrubland. Another gradient of significance can be likened to the tropical gradient running from the dry conditions of hot semi-deserts to savannas or grasslands to woodlands. At the dry extreme an open shrubland occurs (Karroid & Renoster Shrubland, Asteraceous Fynbos), at intermediate positions a herbland occurs (Restioid Fynbos) and at the wet extreme another type of shrubland occurs (Ericaceous Fynbos). This kind of moisture gradient occurs on individual mountains with the xeric end being at the base of the north slopes, and the mesic end being on the upper south aspects. Soil characteristics are closely linked to this gradient; the better developed soils being found on the south aspects. Another moisture gradient is apparent in the vegetation. This gradient runs from the mesic southern coastal mountains to the north west where summer droughts are extremely severe although total rainfall may be similar.Nomenclature follows that used in the Government Herbarium, Stellenbosch.I thank William Bond, Richard Cowling, John Rushworth, Eddy van der Maarel and Marinus Werger for critical comments and discussion.     

Effects of fire on the vegetation of a lowland heathland in North-western Italy

This study focuses on the effect of fire on lowland heathlands at the extreme southern edge of their European distribution (Vauda Nature Reserve, NW Italy). Forty-nine plots (50 m radius) were surveyed between 1999 and 2006. Each year, fire occurrences were recorded and per cent cover of four vegetation types (grassland, heath, low shrubland, and tall shrubland) was estimated in each plot. Vascular plant species richness was also recorded in 255, 1 m² quadrats. After a fire, grassland vegetation expanded, but then declined rapidly as heath and shrubland recovered: 7 years after a fire, tall shrubland encroached on to more than 40% of the plots, and grassland declined from 50% to 20% cover. Between 1999 and 2006, Betula pendula shrubland greatly expanded, while grassland decreased over most of the Reserve, even where fire frequency was high. Tall shrubland had low plant diversity and was dominated by widespread species of lower conservation value. By contrast, early successional vegetation (grassland and low shrubland) had higher richness and more narrowly distributed species, indication that the development of tall shrubland causes significant species loss in the heathland. Italian lowland heathlands are characterized by high rates of shrubland encroachment that threatens both habitat and species diversity. Burning frequencies of once in 3–6 years seem appropriate in this habitat, but burning alone might not suffice without actions to increase herbivore grazing.     

How much evolutionary history in a 10 x 10 m plot?

We use a fully dated phylogenetic tree of the angiosperm families to calculate phylogenetic diversity (PD) in four South African vegetation types with distinct evolutionary histories. Since the branch length values are in this case represented by the ages of plant lineages, PD becomes the cumulative evolutionary age (CEA) of assemblages. Unsurprisingly, total CEA increases with family and with species diversity and observed values are the same as expected from random sampling of family lists. However, when random sampling is done from species lists, observed CEAs are generally lower than expected. In vegetation types which have undergone recent diversification-grassland, fynbos and Nama-karoo-co-occurring species are more closely related than expected, but in subtropical thicket the observed CEAs are well described by random sampling. The use of CEA has great potential for assessing the age of biotic assemblages, particularly as the dating of genus and species-level phylogenies become more accurate.     

Dust as a Nutrient Source for Fynbos Ecosystems,South Africa

Abstract Fynbos is the main vegetation of the Cape Floristic Region, a biodiversity hotspot that occurs in southwestern South Africa. A major question concerning the fynbos ecosystem is how it supports abundant and diverse vegetation on soils derived from nutrient-poor bedrock. In addition to marine aerosols (recycled sea salts), geochemical analyses reported here suggest that dust (aeolian) deposition represents a significant source of nutrients (for example, K, Ca and Zn) to the fynbos ecosystem. Headwater portions of the Boontjies River sub-catchment near the Cederberg Mountains support mountain fynbos communities that are entirely underlain by the Peninsula Formation, a quartz arenite with greater than 98 wt% SiO₂. Fynbos soils in these areas are composed of quartzose sand with 3–6 wt% kaolinitic clay and 1–2 wt% organic carbon. The minor amount of feldspar and mica minerals in the bedrock (0.5 wt% Al₂O₃) suggests an aeolian source for much of the clay minerals in the soil. The isotope composition of soluble Pb and Sr from fynbos vegetation and soils indicates a mixture of anthropogenic and terrigenous sources, most likely from washout of combusted petrol and dust from the arid interior particularly in association with Berg Wind events. Approximate mass balance calculations indicate that washout of aerosols provides an important source of nutrients such as Ca, K, P, Fe, Mn and Zn which the fynbos ecosystem is highly effective in retaining.     

放牧对草地昆虫多样性的影响研究进展

在草地生态系统中,大型草食动物放牧是重要的管理方式之一,对草地生物多样性起着关键的驱动作用。昆虫是草地生态系统中生物多样性的重要组成成分,对生态系统的食物网结构以及其功能与稳定性起着关键作用。已有研究结果表明,大型草食动物与昆虫存在密切联系,放牧对草地昆虫多样性或有正向、或负向、或无明显作用,这依赖于放牧管理方式、昆虫类群以及草地类型。放牧必然通过直接(采食、践踏或粪尿)或间接(植物群落组成或植被结构)作用对昆虫多样性产生显著的影响。当前,关于大型草食动物放牧对草地昆虫多样性影响研究较多,但是,从研究系统性、深入性和延续性来说还存在一定问题。本文在综述国内外对放牧对草地昆虫多样性的影响研究基础上,提出了今后的研究方向,对于理解放牧管理的草地昆虫多样性变化规律,以及为积极探索维持草地昆虫多样性的长期有效的科学管理措施提供理论指导。     

SHORT-TERM RESPONSES OF BIRDS TO FIRE IN OLD MOUNTAIN FYNBOS

FRASER, M. W. 1989. Short-term responses of birds to fire in old Mountain Fynbos. Ostrich: 60:172-182.

Observations of birds were made from six weeks before until one year after a fire in 28-year-old Mountain Fynbos vegetation near Stellenbosch, Cape Province, South Africa. No fire attendance by birds was observed, in contrast to observations made at fires in other southern African vegetation types. Bird species composition in the study site as a whole was virtually unchanged after the fire. With the exception of the density of nectarivorous birds (which decreased), variation in the bird densities of study plots in six vegetation structural formations was considered to be as much seasonal as fire-related.     

Shrubland Restoration Following Woody Alien Invasion and Mining: Effects of Topsoil Depth, Seed Source, and Fertilizer Addition

Invasion by woody alien plants, construction, and mining operations are among the major disturbances degrading vegetation in the Cape Floristic Kingdom, South Africa. The aim of this study was to assess whether native fynbos shrubland vegetation could be restored following dense alien invasion and disturbance by mining. An area supporting dense alien trees was cleared and topsoil was stripped and stockpiled to simulate mining disturbance. A field trial investigated the effects of topsoil depth, seed mix application, and fertilizer on native species recruitment and vegetation development over a three‐year period. Soil‐stored seed banks contributed 60% of the species recruited, indicating that areas invaded for three decades have good restoration potential. The addition of a fynbos seed mix, which included serotinous overstory species, improved both the richness and structural composition of the vegetation. Most species sown in untopsoiled plots established, but survival and growth was low compared to topsoil plots. Poor growth in combination with a lack of soil seed bank species, indicate that restoring a diverse and functional cover of indigenous vegetation on subsoil is not possible in the short‐term. Soil amelioration is required to improve rooting conditions and initiate ecosystem processes. Shallow and deep topsoil treatments yielded high plant density, richness, and projected canopy cover, but canopy cover was higher in deep topsoil plots throughout the trial. Fertilizer addition increased canopy cover in untopsoiled and shallow topsoil plots via an increase in alien annual species. Fertilizer addition ultimately may lead to increased native vegetation cover in untopsoiled areas, but as it increased proteoid mortality on deep topsoil plots, it is not recommended for sites where topsoil is available. A species‐rich and structurally representative fynbos community may be restored on topsoiled areas provided that the native disturbance regime is simulated and seeds of major structural guilds not present in the soil seed bank are included in the seed mix.     

Island-dominated landscapes of large floodplain rivers, a European perspective

1. The landscapes of large floodplain rivers are characterised by heterogeneous environments related to the interplay of flood flows, sediment transport and vegetation dynamics.
2. The large rivers of Europe, and probably most rivers throughout the forest biomes, were characterised by islands but over the period of major human interference, many have become dominated by incision and narrowing so that they are now characterised by single-thread and relatively simple channel forms.
3. Vegetation plays an active role in developing heterogeneous channel forms through (a) biotic processes such as seed dispersal, vegetative regeneration and succession and (b) abiotic effects such as increasing flow resistance inducing sedimentation, and decreasing bank erodibility.
4. In particular, accumulations of living driftwood (cf. dead driftwood accumulations and dispersed seedlings) accelerate sedimentation and island development.
5. River reaches with vegetated islands have a high habitat diversity.
6. The natural influences of flood disturbance, wood accumulation, vegetation growth, island development and tree die-off, cause island-dominated reaches to undergo cycles of island growth and decay that are related to cycles of aquatic habitat diversification and simplification.     

Diversity,composition and guild structure relationships between soil-stored seed banks and mature vegetation in alien plant-invaded South African fynbos shrublands

We investigated vegetation-seed bank relationships at three fynbos sites on the Cape Peninsula, South Africa, and the impacts to these sites of invasion by the alien tree Acacia saligna. Soil-stored seed banks in uninvaded fynbos were of a similar density to those previously measured in fynbos (ca. 1100–1500 seeds m^-2) and were dominated by mostly short-lived species. Lack of similarity between mature vegetation and seed banks, suggests that seed banks are poor predictors of mature vegetation composition and structure in fynbos. This lack of correspondence was attributed to the ephemerals (present only in the soil seed bank) and the dominance of serotinous (aerial seed bank) and sprouting (soil seed bank low to absent) species, in mature vegetation. Long-lived seeders were among the 10 most abundant species in the seed banks at all sites and at two sites shrub species contributed more to seed bank richness than any other growth form. Soil-stored seed banks, therefore, boost species richness and diversity both in early post-fire and later seral stages.There was a decline in fynbos species richness, diversity and abundance both in the standing vegetation and seed banks with increasing duration of invasion by the alien tree, Acacia saligna. However, the rate of decline was higher for the vegetation than the seed banks, suggesting that many fynbos species have long-term persistent seed banks. At two sites, there was no obvious shift in community composition associated with Acacia invasion: invaded sites were depauperate versions of the uninvaded site. However, at a third site, the vegetation composition shifted towards a community dominated by bird-dispersed thicket species and its seed bank shifted towards a community dominated by wind-dispersed perennials. Community composition of the soil seed banks under dense, recent Acacia was very similar to that of the corresponding uninvaded fynbos at all sites, indicating that there is good potential to return to species-rich fynbos vegetation after removal of the alien Acacia. Most seed bank species persisted in the soil seed bank of the long-invaded fynbos at low frequency and density, indicating high seed longevity in many species. We suggest that either a thick Acacia litter layer or a deep (>5 cm) burial moderated the fire and ambient temperature effects, preventing these seeds from germinating after fire and thus preventing loss from the seed bank.     

Microbial diversity and community structure in Fynbos soil

The Fynbos biome in South Africa is renowned for its high plant diversity and the conservation of this area is particularly important for the region. This is especially true in the case of endangered vegetation types on the lowlands such as Sand Fynbos, of which only small fragments remain. The question is thus whether the diversity of the above‐ground flora is mirrored in the below‐ground microbial communities. In order to determine the relationship of the above‐ and below‐ground communities, the soil community composition of both fungal and bacterial groups in Sand Fynbos was characterized over space and time. A molecular approach was used based on the isolation of total soil genomic DNA and automated ribosomal intergenic spacer analysis of bacterial and fungal communities. Soil from four different sites was compared to resolve the microbial diversity of eubacterial and fungal groups on a local (alpha diversity) scale as well as a landscape scale (beta diversity). The community structures from different sites were compared and found to exhibit strong spatial patterns which remained stable over time. The plant community data were compared with the fungal and the bacterial communities. We concluded that the microbial communities in the Sand Fynbos are highly diverse and closely linked to the above‐ground floral communities.