Response to nutrient additions by the plant growth forms of sand-plain lowland fynbos,South Africa

The growth response (foliage projective cover: FPC) of the plant growth forms of lowland fynbos, South Africa to a complete factorial fertilizer addition of nitrogen (N), phosphorus (P) and a mixture of all essential nutrients excluding N and P (M) was monitored for two years. Ordination by correspondence analysis revealed a successional trend as the vegetation aged, with increases in proteoid, and to a lesser extent restioid, and decreases in reseeding ericoid, graminoid, geophyte and annual plant FPC but no discernible effects due to fertilizer application. Analysis of covariance revealed significant increases in restioid, graminoid and annual plant FPC with N-addition, 1 and 2 yr after fertilizer addition and of total FPC with N-addition for only one year. Of the other nutrient treatments, only an increase in annuals with P-addition and a reduction in the rate of decline of reseeding ericoid with M-addition were found, both after two years. Thus nitrogen may often limit vegetative growth of at least the herbaceous species. The herbaceous growth forms are more plastic in their morphological growth responses to nutrient additions than the slow-growing, stress tolerant, evergreen shrub species. The vegetation appears to be resilient, at least in the short-term, to a disturbance of this magnitude. However, chronic nutrient applications and those of larger magnitude will probably result in long-term changes in species composition, with an increase in ephemeral, nutrient demanding species.Abbreviations FPC = Foliage Projective Cover     

Dieback and mortality of South African fynbos shrubs is likely driven by a novel pathogen and pathogen‐induced hydraulic failure

We examined whether extensive dry season dieback and mortality in a South African fynbos community were due to drought or pathogen attack. Plant dieback and mortality have been reported elsewhere in similar plant communities suggesting potential for a widespread climatic or biotic threat to this community. We collected tissue samples from Brunia noduliflora, the dominant plant in the community, and cultured them for potential plant pathogens. We also measured dry season predawn and midday water potentials of healthy and stressed plants and constructed pressure‐volume curves to assess turgor loss point. Plant stress and mortality were monitored over a 2‐year study period. Both healthy plants and plants that displayed moderate signs of stress had dry season predawn water potentials well above their turgor loss point suggesting plants were not water stressed. However, plants displaying >60% crown dieback had much lower water potentials (as low as ?12 MPa). A previously undescribed fungus (Pythium sp.) was isolated from the root vascular tissue of all stressed plants and was not present in healthy plants. The proximate cause of plant stress was likely pathogen‐induced, while the ultimate cause of plant death appears to be extreme water stress. The present study suggests that Brunia (Bruniceae), Leucadendron (Proteaceae) and Erica (Ericaceae), all emblematic and dominant genera within the diverse fynbos community, may be susceptible to Pythium infection. This may pose a serious threat to communities already threatened by climate change.     

Winter growth phenology and leaf orientation in Pachypodium namaquanum (Apocynaceae) in the succulent karoo of the Richtersveld,South Africa

Pachypodium namaquanum (Nyley ex Harb.) Welw., an unusual arborescent stem succulent from the succulent karoo of the arid Richtersveld in north-western South Africa and adjacent Namibia, is characterized by a striking curvature of the terminal 20–60 cm of the trunk toward the north. This orientation displays the single terminal whorl of drought-deciduous leaves with their flat surface angled at a mean inclination of 55° from horizontal. Inclination of 50–60° was found in 65% of individuals sampled, and 85% were inclined between 45 and 65°. Northward azimuth was also quite regular, but varied slightly between populations. The fixed leaf orientation in P. namaquanum maximizes radiation absorption during the winter months when leaves are present. Leaves normally form in early fall (April) and abscise early in spring (October). Growing season conditions in the Richtersveld are relatively mild, with mean maximum temperature dropping only to 21.6°C in July, the coldest month of the year. Frosts are rare. By the fixed orientation of its leaf whorl, P. namaquanum is able to maintain nearly twice the midwinter radiation absorptance that it would have with horizontal orientation. Over an annual cycle the angled leaves receive more radiation than would horizontal leaves for each of the 6 months in which they are present on the plant. This increased winter irradiance is hypothesized to singificantly increase net primary production by concentrating growth activities in winter months and allowing the species to remain dormant during the hyperarid conditions of the hot summer months. Midwinter flowering from apical buds in P. namaquanum may also be aided by its stem orientation. The evolution of this characteristic pattern of winter growth phenology and nodding stem orientation may have come about because of low but relatively regular autumn precipitation and moderate winter temperatures. Slow and regular growth of P. namaquanum leads to long lifespans which may reach 300 years or more.     

Demography of the striped mouse (Rhabdomys pumilio) in the succulent karoo

The striped mouse (Rhabdomys pumilio) is widely distributed in southern Africa, inhabiting a wide range of habitats. We describe the demography of the striped mouse in the arid succulent karoo of South Africa, and compare our findings with those of published results for the same species from the moist grasslands of South Africa. In both habitats, breeding starts in spring, but the breeding season in the succulent karoo is only half as long as in the grasslands, which can be explained by different patterns and levels of rainfall; the succulent karoo receives mainly winter rain and rainfall is much less (about 160 mm year⁻¹) than in the grasslands (>1000 mm year⁻¹) which experience summer rain. Population density increased from 37 (start of breeding season) to 171 (end of breeding season) mice per hectare. A high yearly survival rate of 27% during our study from summer to the next breeding season resulted in a population density that was 10 times higher in the succulent karoo than in grasslands. The comparatively high population density may result in habitat saturation and thus forced philopatry, promoting group living in the succulent karoo, which contrasts with the solitary life-style exhibited by populations in moist grasslands.

Zusammenfassung

Demographie der Striemengrasmaus (Rhabdomys pumilio) in der SukkulentenkarooDie Striemengrasmaus (Rhabdomys pumilio) ist im südlichen Afrika weit verbreitet und kommt in unterschiedlichen Habitaten vor. In dieser Studie beschreiben wir die Demographie der Striemengrasmaus in der ariden Sukkulentenkaroo Südafrikas, und vergleichen unsere Ergebnisse mit den Ergebnissen früherer Studien, welche dieselbe Art in den feuchten Grasländern Südafrikas untersucht haben. Die Fortpflanzungssaison beginnt in beiden Habitaten im Frühling, aber sie ist in der Sukkulentenkaroo nur halb so lange wie in den Grasländern. Der Grund dafür wird im unterschiedlichen Niederschlagsmuster vermutet: In der Sukkulentenkaroo regnet es wenig im Winter (ca. 160 mm), während es in den Grasländern viel im Sommer regnet (>1000 mm). Die Überlebensrate in der Sukkulentenkaroo war vom Sommer zur nächsten Fortpflanzungssaison im Frühjahr mit 27% außerordentlich hoch, während die Populationsdichte zehnmal so groß war wie in den Grasländern. Am Anfang der Fortpflanzungssaison betrug die Populationsdichte 37 Mäuse ha⁻¹, am Ende der Fortpflanzungssaison 171 Mäuse ha⁻¹. Die außergewöhnlich hohe Populationsdichte führt womöglich zu Habitatsättigung und einem Fehlen freier Territorien, was mit erklären könnte, warum die Striemengrasmaus in der Sukkulentenkaroo gruppenlebend ist, in den Grasländern hingegen solitär.     

Beating the blaze: Fire survival in the fan aloe (Kumara plicatilis), a succulent monocotyledonous tree endemic to the Cape fynbos,South Africa

Fire is central to the ecology of Mediterranean‐type climate ecosystems, but little is known about the fire ecology of succulent plants therein. This study investigated the fire ecology of an arborescent succulent monocot, Kumara plicatilis (L.) G. D. Rowley (Asphodelaceae), a Cape fynbos endemic. Habitat suitability was assessed to determine whether the species tolerates or ‘avoids’ fire, and fire survival traits (bark thickness and tissue water content) were measured. The population size structure and density of three K. plicatilis populations were assessed after natural fires, and resprouting potential was investigated. Kumara plicatilis adopts a dual fire survival strategy, occupying rocky sites to ‘avoid’ fire and possessing morphological features that afford fire tolerance, e.g. well‐protected apical meristems and thick corky bark. Bark thickness of burned individuals in situ was similar to unburned plants, suggesting that K. plicatilis bark provides effective insulation against fire. Mortality rates were 64%, 40% and 11%, and decreased as rock cover at the population level increased. All three populations showed reduced plant density post‐fire, with greater density reductions associated with lower rock cover. Small plants appear most vulnerable to fire damage due to lower absolute bark thickness and plant heights within the flame zone. Kumara plicatilis is an apical sprouter, recovering after fire or mechanical stem damage by onward growth from surviving stem apices, rather than resprouting. Post‐fire population recovery therefore likely depends on inter‐fire recruitment.     

Abiotic and biotic determinants of the seasonal dynamics of the tick Rhipicephalus appendiculatus in South Africa

Abstract. The creation of a generic population model for the tick Rhipicephalus appendiculatus requires a detailed, quantified understanding of the interactions of these ticks with their biotic and abiotic environment in the different parts of their range, from the tropical regions of East Africa to the temperate regions of South Africa. The much greater seasonal variation in climatic conditions, particularly temperature, further from the equator introduces variable development rates and diapause into the life cycle. Estimates of natural temperature-dependent interstadial development periods, derived from a combination of published laboratory and field data, were applied to published data on the seasonal abundance of R.appendiculatus on three farms in the Eastern Cape Province of South Africa. This enabled an assessment of which ticks of one stage give rise to which ticks of the next stage, from which (a) the onset and duration of diapause in unfed adults, and (b) seasonal interstadial mortality indices in the form of fc-values, could be estimated. The contribution of biotic (tick density) and abiotic (climatic) factors as predictors of mortality at each life stage was investigated by step-wise multiple regression. Density-independent mortality at the female-to-larval stage is correlated with geographically variable climatic factors, minimum temperature at two farms and minimum relative humidity at the third. The other two stages are governed by density-dependent mortality, which, it is argued, may be caused largely by the hosts' acquired resistance to ticks. As expected on theoretical grounds, this density dependence is weaker nearer to the edge of the tick's range and for the more vulnerable immature stages.     

Root of edaphically controlled Proteaceae turnover on the Agulhas Plain, South Africa: phosphate uptake regulation and growth

The influence of phosphorus (P) availability on growth and P uptake was investigated in South African Proteaceae: (1) Protea compacta R.Br., endemic on severely nutrient-impoverished colluvial sands; (2) Protea obtusifolia Bueck ex Meissner; and (3) Leucadendron meridianum I. J. Williams, the latter both endemic on comparatively fertile limestone-derived soils. Plants were grown hydroponically in 1000 L tanks at 0.01, 0.1 or 1.0 microm P for 14 weeks. Biomass accumulation was influenced by P availability, doubling as [P] increased from 0.1 to 1.0 microm. Total biomass was greatest for P. compacta, but L. meridianum and P. obtusifolia had two to four times greater relative biomass accumulation at 0.1 and 1.0 microm [P]. Proteoid root clusters developed at both 0.01 and 0.1 microm[P], but were suppressed at 1.0 microm [P]; this was a 10-fold lower [P] than previously reported to inhibit cluster root formation. Rates of net P uptake at 5 microm P decreased in response to increased P availability from 0.01 to 1.0 microm P. Significant between-species differences in rates of P uptake and capacity to down-regulate P uptake were observed: P. compacta < P. obtusifolia < L. meridianum. The species responses are discussed in terms of adaptation to mosaics in soil P availability and the high beta diversity in the natural habitat.     

Reproductive ecology of Aloe plicatilis,a fynbos tree aloe endemic to the Cape Winelands,South Africa

While the pollination ecology of many Aloe species is well-documented, knowledge on aloe seed ecology, and hence aloe reproductive ecology in its entirety is limited. The aim of this study was to investigate the reproductive ecology of Aloe plicatilis, a Cape fynbos tree aloe endemic to the Cape Winelands, South Africa. Results from a pollinator exclusion experiment conducted at an A. plicatilis population on Paarl mountain suggests pollination primarily by insects, although bird visitation significantly increased seed set/fruit indicating possible co-pollination with insects. The species' long-tubed flowers and production of concentrated nectar, with observations of malachite sunbirds as the most common avian visitors to A. plicatilis flowers indicate the importance of long-billed specialist avian nectarivores as floral visitors. Analysis of the relationship between plant size and inflorescence production for five populations combined revealed a significant, positive linear relationship between plant size and the logarithm of the number of inflorescences/plant. Natural fruit and seed set determined for three populations (1325, 27,930 and 251,616 seeds/population) suggests low reproductive output compared to several other Aloe species. The smallest (31 individuals) and least dense (75 plants/ha) A. plicatilis population produced the lowest seed set/plant (128 seeds) and per population (1325 seeds), suggesting an Allee effect. Evaluation of seed dispersal potential showed that potential dispersal distances were approximately three times the canopy height; however, the occurrence of A. plicatilis on mountains isolated from more continuous mountain ranges on which the species also occurs suggests the possibility of long-distance dispersal by strong, gusty, summer winds. Soil seed bank samples collected from 13 populations yielded close to zero seedling emergence, indicating the absence of persistent seed banks. A. plicatilis seeds stored under ambient laboratory conditions for 3, 18 and 24 months were germinated in an environmental control chamber and a laboratory. High percentage germination was recorded for 18- and 24-month-seed (86 and 80%, respectively), while germination of 3-month-old seeds was three times lower, suggesting the need for after-ripening. Germination of fresh and one-year-old seed under ambient nursery conditions at the Karoo Desert National Botanical Garden in Worcester yielded emergence percentages of 67 and 44%, respectively, and were therefore less successful than germination under more controlled conditions. This is the first known study to investigate the reproductive ecology of a tree aloe species and that of a Cape fynbos aloe. The study highlights the importance of further studies on aloe seed ecology, particularly for rare and threatened species.     

Conservation biology of the succulent shrub,Euphorbia barnardii,a serpentine endemic of the Northern Province,South Africa

Euphorbia barnardii White, Dyer & Sloane is a relatively small, succulent shrub found in the Northern Province of South Africa. In 1994 it was listed as endangered by the Transvaal Threatened Plants Programme, because only three populations, totalling 1150 plants, were found. However, our study found five populations totalling 10 783 plants (9503 were reproductive). One population, found 50 km from the others, in the Bewaarkloof district, differs in terms of habitat type and morphology and may therefore be a different taxon. The other populations occur in Sekhukhuneland. Soil analyses showed that E. barnardii is restricted to ultramafic (serpentine) substrates. A sample of 2015 plants was examined in 1995 for size, stage, new growth, dead branches, reproduction (indices of vigour), damage and disease. Analysis of the results of 10 years’ demographic monitoring (1985–1995) on two of the Sekhukhuneland populations showed that one population, which was vigorous in 1995, showed little change in population size and structure, while the other, which had low vigour in 1995, showed a precipitous decline to local extinction in the monitoring plot. Aerial photographs taken in 1957 (1963 for Bewaarkloof) and 1986 showed large increases in human population density within 1.5 km of the E. barnardii populations except at Bewaarkloof, where it decreased. An increase in human habitation is associated with an increase in livestock (mostly cattle and goats) which trample plants. Trampling damages the terminal segments, which may lead to lowered reproductive output and increase susceptibility to opportunistic bacterial wilt pathogens. Bacterial wilts were prevalent on all populations but were particularly high at Bewaarkloof and on the population that showed a steep decline in numbers. Disease incidence was associated with the level of plant damage. The impact on plants further up the slopes and on the crest of hills (quite far from human settlements) was to a far lesser degree but further increases in human population density could change this situation. While this study has shown that the population size of E. barnardii is much greater than previously thought, the species is still threatened by several different processes and should be listed as ‘Vulnerable (A1a + c, B1, B2b + e, C1, D2)’ according to World Conservation Union categories.     

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.     

Remarkable medium-term dynamics of leaf succulent Mesembryanthemaceae shrubs in the winter-rainfall desert of northwestern Namaqualand,South Africa

Populations of shrubs in a winter-rainfall (ca. 70 mm yr_-1 desert community (Succulent Karoo), dominated by leaf succulents, showed remarkable dynamics over a 17-yr period. After a severe drought in 1979, which caused high plant mortality, perennial species number in a permanent 10 m×10 m plot doubled between 1980 and 1996, when the maximum of 41 species was recorded. Numbers of individuals of evergreen, leaf succulent shrubs also doubled over the same period, but showed fluctuations in response to dry years during the monitoring period. Detailed observations on the four dominant leaf succulent shrubs (all members of the Mesembryanthemaceae) between 1983 and 1996, showed species-specific patterns in population turnover. Mortality of all observed individuals ranged between 60% and 85%, and the proportion of the population that was recruited over this period ranged from 62% to 89%. In only one species did individuals persist throughout the monitoring period. Mean ages of individuals, excluding seedlings with lifespans of <1 yr, and individuals observed at the beginning of the monitoring period, ranged from 4.6 yr to 5.6 yr. Patterns of mortality and recruitment showed substantial differences among species and were not all attributable to rainfall patterns. Overall, the turnover of the shrub populations over the 15-yr monitoring period was remarkably high for a system of desert perennials. These unusual population patterns may explain the unique structure of leaf succulent-dominated communities in the Succulent Karoo.     

A large male hominin cranium from Sterkfontein, South Africa, and the status ofAustralopithecus africanus

Stw 505 is the most complete hominin cranium discovered in Sterkfontein Member 4 since Broom's excavations. It was found in situ in Member 4 breccia in 1989 and is larger, on the whole, than any other cranium from Sterkfontein that has comparable parts. Displacement due to breakage, as well as plastic deformation, has affected Stw 505 in several areas, especially the face and the vault. Diagnosticmorphology is nevertheless abundant in the specimen. In several areas-the distinct anterior pillar, the straight inferior border of the zygoma, the pattern of cresting on the naso-alveolar clivus, the basal aspect of the temporal bone-Stw 505 closely matches the morphology of specimens of Australopithecus africanus and is distinct from other hominins. Some isolated characters overlap with other groups, mainly early Homo and/or A. robustus. However, only the hypodigm of A. africanus can accommodate the entire suite of morphology.In some cases, Stw 505 introduces more variation into the Sterkfontein sample. For example, prominent superciliary eminences occupy the medial portions of the supraorbital region and flow medially into a strongly protruding glabellar mound. These characteristics are probably attributable to sexual dimorphism. In many respects, Stw 505 highlights similarities between A. africanus and early Homo. Comparison with other species suggests that males of A. africanus do not show derived features of A. robustus that are not also present in females, and that cranial differences between A. afarensis and A. africanus have, if anything, been understated.     

Physiological responses to folivory and phytopathogens in a riparian tree,Brabejum stellatifolium,native to the fynbos biome of South Africa

The canopies of many tree species sustain a large diversity of folivorous arthropods and phytopathogenic fungi. These organisms are thought to influence overall tree and stand productivity. Leaf diseases caused by Phyllosticta owaniana and Periconiella velutina, phytopathogenic fungi commonly found on the native riparian tree Brabejum stellatifolium (wild almond), like any other leaf disease, can potentially reduce a plant's photosynthetic efficiency. In addition to these two phytopathogens, the weevils Setapion provinciale and Setapion quantillum are abundant in wild almond canopies. Despite their pervasive occurrence, the impacts of these phytopathogens and arthropods on host tree leaf physiology have not been examined. The gas exchange response of wild almond leaves to phytopathogens and folivore damage was assessed. Leaf nitrogen, phosphorus and water content were also determined. Declines in photosynthetic rates and other physiological parameters were associated with increasing damage severity by weevils and phytopathogens in leaves of B. stellatifolium. Nitrogen and phosphorus contents were negatively associated with disease severity. Water and phosphorus contents were also negatively correlated with increased weevil damage, while nitrogen content was positively correlated with it. The observed responses of B. stellatifolium metabolic functioning to fungal pathogen and folivory indicate a possibility of suppressed wild populations of wild almond.