Ecological and phylogenetic patterns of carbon isotope discrimination in the winter-rainfall flora of the Richtersveld,South Africa

Leaf tissues of vascular plant species from the arid Richtersveld of northern Namaqualand (Succulent Karoo), South Africa, show a great range of variation in carbon isotope discrimination. Analyses of values for species with obligate C₃ metabolism demonstrated that the water use efficiency indicated by these values varies between species and within a species population both between seedlings and mature shrubs and between sites along an aridity gradient. An expected pattern of increasing water use efficiency with greater longevity in these species was not present, however. Only among ephemerals were there taxa with high values associated with low water use efficiency, but these high values were balanced by other ephemeral species with low values comparable to those typical of moderate to long-lived shrubs. The extent to which putative CAM species in the succulent flora rely entirely on nocturnal carbon fixation is highly variable. Species of Crassulaceae and Aloe (Asphodelaceae) relied almost exclusively on CAM photosynthesis, although Crassulaceae from more mesic environments may be CAM-flexible. Succulent-leaved species of Senecio and leaves of stem-succulent Euphorbia were CAM-flexible in their values, while shrubby species of Ceraria (Portulacaceae) appeared to rely primarily on C₃ photosynthesis. Variable patterns of reliance on CAM photosynthesis were present in the ecologically dominant Mesembryanthema of the Aizoaceae. Shrubby species of the subfamily Mesembryanthemoideae, although capable of CAM-flexibility in less xeric habitats, relied largely on CAM fixation of carbon in our field sites. Within the subfamily Ruschioideae, there was a wide range of values indicating species with CAM, CAM flexibility, and primary reliance on C₃ photosynthesis. Low values indicative of typical CAM fixation were associated with species with greater longevity, suggesting that a positive correlation may exist between the degree of reliance on CAM photosynthesis and plant life span.     

Sex differences in obesity rates in poor countries: Evidence from South Africa

Globally, men and women face markedly different risks of obesity. In all but of handful of (primarily Western European) countries, obesity is much more prevalent among women than men. We examine several potential explanations for this phenomenon. We analyze differences between men and women in reports and effects of potential underlying causes of obesity—childhood and adult poverty, depression, and attitudes about obesity. We evaluate the evidence for each explanation using data collected in an urban African township in the Cape Town metropolitan area. Three factors explain the greater obesity rates we find among women. Women who were nutritionally deprived as children are significantly more likely to be obese as adults, while men who were deprived as children face no greater risk. In addition, women of higher adult socioeconomic status are significantly more likely to be obese, which is not true for men. These two factors - childhood circumstances and adult SES - can fully explain the difference in obesity rates between men and women that we find in our sample. More speculatively, in South Africa, women's perceptions of an ‘ideal’ female body are larger than men's perceptions of the ‘ideal’ male body, and individuals with larger ‘ideal’ body images are significantly more likely to be obese.     

Geographical affinities of the Cape flora, South Africa

Aim The flora characteristic of the Cape Floristic Region (CFR) is dominated by a relatively small number of clades that have been proposed as ‘Cape clades’. These clades have variously been suggested to have African or Austral affinities. Here we evaluate the support for these conflicting hypotheses. In addition, we test the hypothesis that these clades share a common time of differentiation from their geographical neighbours. Location The Cape Floristic Region, South Africa Methods We use both published and unpublished phylogenetic information to investigate the geographical sister areas of the Cape clades as well as the timing and the direction of biogeographical disjunctions. Results Almost half of the Cape clades for which unambiguous sister areas could be established show a trans‐Indian Ocean disjunction. The earliest trans‐Indian Ocean disjunction dates from 80 Ma. Other disjunctions date from various times in the Cenozoic, and we suggest that the process of recruiting lineages into the Cape flora might be ongoing. Relatively few Cape clades show a sister relationship with South America and tropical Africa, despite their relative geographical proximity. Numerous Cape clades contain species also found on tropical African mountains; in all cases tested, these species are shown to be embedded within the Cape clades. While many Cape clades show a relationship with the Eurasian temperate flora, this is complicated by their presence in tropical Africa. The single case study addressing this to date suggests that the Cape clade is nested within a European grade. Main conclusions Although many Cape clades show Austral rather than African relationships, there are numerous other patterns suggestive of a cosmopolitan flora. This spatial variation is echoed in the temporal data, from which, although there is wide variance around the dates of disjunctions, it is clear the Cape flora has been assembled over a long time period. There is no simple hypothesis that can account for the geographical sources of the currently distinctive Cape flora. The phylogenetic positions of Afromontane members of Cape clades suggest a history of dispersal from the CFR, rather than the reverse.     

Evaluation of Restoration Techniques for the Succulent Karoo, South Africa

Abstract Possible constraints on the passive recovery of bare areas in the Karoo, a semiarid region in South Africa, include inadequate supply of seed, availability of suitable microsites for plant establishment, altered soil properties, and the truncation of key soil biotic processes. Here we investigate the possibility of initiating the restoration of bare areas by soil surface treatments with gypsum (CaSO₄) and/or organic mulch. We also apply an exogenous seed source to test the hypothesis that seed availability limits autogenic recovery. Both gypsum and mulch improved rain water infiltration, gypsum more so than mulch, and both treatments resulted in significantly higher numbers of reseeded seedlings compared with controls. Gypsum also improved the survival of the cohorts of seedlings of the larger seeded Tripteris sinuata. Tripteris showed the highest number of seedlings (maximum count of 150 seedlings/1,000 viable seeds sown) and surviving plants of the three reseeded species, which included two small‐seeded species, Ruschia spinosa and Chaetobromus dregeanus. Throughout the study period significantly higher plant volumes of naturally seeded annuals and perennials were recorded in the gypsum and/or mulch treatments compared with the controls. Germination and emergence of reseeded and naturally seeded plants appears to be determined by the availability of cool season (autumn to spring ) soil moisture, whereas follow‐up rainfall during this time is important for plant survival. Mulching of bare areas in the Succulent Karoo has the potential to re‐create vegetated areas that will further capture and conserve water, soil, and nutrients. Gypsum also showed positive results but might not be a cost‐effective option because of transport costs to these remote arid areas.     

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.     

A contribution to the rust flora (Uredinales) of southern Africa,with an emphasis on Namibia

Five new rust species are described and hitherto unknown spore states for the following seven species are reported: Puccinia desertorum on Evolvulus alsinoides, Uromyces comptus on Merremia bipinnatipartita, Puccinia halsei on Acacia hereroensis, Ravenelia transvaalensis on Acacia mellifera, Puccinia abutili on Abutilon angulatum, on Abutilon cf. austroafricanum, and on Abutilon cf. rehmannii, Puccinia lycii on Lycium sp. and Puccinia turgida on Lycium europaeum and on Lycium cf. oxycarpum. We also examined Uredo combreticola on Combretum cf. engleri, on Combretum hereroense, and on Combretum zeyheri, Puccinia afra on Lycium sp., and Uredopeltis cf. chevalieri on Grewia flavescens. All mentioned rust fungi are described in detail and are shown by line drawings. Selected species are illustrated with SEM-photographs. Part 220 in the series “Studies in Heterobasidiomycetes” from the Botanical Institute, University of Tübingen     

Biodiversity,habitat and range-size aspects of a flora from a winter-rainfall desert in north-western Namaqualand,South Africa

We examined patterns of richness and endemism in relation to phylogeny (family membership), growth form and habitat in the flora of an arid (<60 mm annual rainfall), sandy coastal belt between Port Nolloth and Alexander Bay on the north-west coast of Namaqualand, South Africa. This region forms part of the species-rich Succulent Karoo biome. In an area of 750 km₂, 300 plant species were recorded with 24% endemnicity. This level of diversity and endemism is exceptionally high for a desert region. Plant distribution is strongly influenced by habitat type. Diversity, especially of endemic species, was concentrated on rocky or unusual (lag-gravel pavements) habitats. Endemics were not random phylogenetic nor biological assemblages. Dwarf leaf-succulent shrubs, especially Mesembryanthemaceae, were significantly over-represented amongst the endemic component. The results are discussed in terms of a model for plant diversification in an arid environment with relatively predictable sources of moisture.     

The Potential of Cephalophyllum inaequale (L. Bolus) for the Restoration of Degraded Arid Landscapes in Namaqualand, South Africa

The potential of Cephalophyllum inaequale was investigated for use in initiating ecosystem restoration in degraded landscapes of Namaqualand, South Africa. Cephalophyllum inaequale, a perennial shrub, is a member of the succulent Mesembryanthemaceae family, typical of the Succulent Karoo Biome and in particular of the Namaqualand area. A bioclimatic envelope was modeled to establish the area in which this species might feasibly be used. The regional bioclimatic potential for C. inaequale proved to be extensive, covering approximately 17,500 km². An examination of the functional role of C. inaequale showed it to facilitate early seedling survival in this community. A nearest‐neighbor study found no evidence of interspecific competition between C. inaequale and its dominant co‐occurring species, possibly due to vertical stratification of rooting structures. Cephalophyllum inaequale significantly reduced wind speed and soil erosion. Experiments to test the feasibility of propagating, reintroducing, and establishing this species showed that it easily germinates from seed, and transplanted cuttings have a high survival rate. This study demonstrates that C. inaequale has potential for use in initiating the restoration of degraded lands in South Africa .     

Status of succulent shrubs in the southern Namib Desert, Succulent Karoo Biome

The ratio of dead to alive succulent shrubs as an indicator of turnover was investigated to test whether the accepted notion of rapid turnover rates and cyclic succession in the Succulent Karoo Biome are applicable in the southern Namib. Based on counts of dead and alive plants, twelve species in two habitat types were investigated. These short‐term data which could not incorporate recruitment rates or changes over time, generated, however, some hypotheses. (i) Rapid turnover rates are likely not supported by all succulent plants at the Succulent Karoo Biome's northern boundary. With the exception of one species, mortality rates in succulent shrubs across a range of plant functional types were lower than reported in other parts of the biome. (ii) Sand plains appeared to support some transient species, indicating that there may be differences in vegetation dynamics between habitats. (iii) Species of different plant functional types showed no differences in ratio of dead to alive plants related to habitat. These species may have broad ecological tolerance limits and are perhaps less affected by changes in their environment. (iv) Shrubby Mesembryanthemaceae have high turnover rates and hence a short live span in the investigated area, but are longer lived than elsewhere in the Succulent Karoo Biome. As these hypotheses have implications for management and conservation of succulent species in this global biodiversity hotspot, a detailed evaluation of vegetation turnover, balancing mortality versus recruitment, should be investigated over a longer time span.     

Biogeography of prostrate-leaved geophytes in semi-arid South Africa: hypotheses on functionality

Nowhere is the species diversity of geophytes greater than in the five mediterranean-climate ecosystems of the world. Of these, the Cape mediterranean zone of South Africa is the most speciose. While the relative diversity and importance of geophytes of all of the other four mediterranean regions of the world drops off sharply as one moves into adjacent winter-rainfall desert regions, geophytes in the semi-arid to arid Succulent Karoo (including Namaqualand) remain a very important component of the flora, both in terms of abundance and diversity (comprising 13 to 29% of the regional floras in this region). Apart from species richness, there are also a number of interesting geophyte growth forms in this region. One unusual growth form is geophytes with flattened leaves that lie prostrate on the soil surface. At least eight families (Amaryllidaceae, Colchicaceae, Eriospermaceae, Geraniaceae, Hyacinthaceae, Iridaceae, Orchidaceae and Oxalidaceae) exhibit this growth form. While this growth form is relatively common in many geophyte lineages in the Succulent Karoo biome and the Cape mediterranean zone (Fynbos biome), and occurs infrequently through the summer-rainfall temperate regions of Africa, it is virtually absent in other regions worldwide. A null hypothesis is that the prostrate leaved trait is a neutral characteristic, however biogeographical data do not support this. A neutral trait would be unlikely to show such a clear pattern of distribution. Several alternative hypotheses on the adaptive significance of this growth form are discussed. These include: avoidance of herbivory, reduction in competition from neighbors, creation of a CO₂ enriched environment below the leaves, reduction of water loss around the roots, reduction of water loss through transpiration, precipitation of dew on the leaves and maintenance of optimal leaf temperatures for growth.     

The biocultural evolution of Khoisan populations of Southern Africa

The research presented here offers new information on the recent evolution of Khoisan populations of southern Africa through the new study of Holocene skeletons. When combined with subfossil and historical remains, these archeological specimens provide a skeletal record for the last 5000–9000 years of southern African prehistory. Multivariate statistical analyses of cranial measurements were used to determine patterns of morphological variation in the skeletal record with which hypotheses of biocultural evolution were tested. These analyses yielded the following results. First, the traditional distinction between Bushmen and Hottentots holds for recent inland individuals. Second, there is a suggestion of a morphologically distinct San population living on the coast of South Africa. The idea of a “Strandloper” population suggested by early scholars is here revived. The third result is that there is little evidence of complete population succession on the coast of South Africa, suggesting that there were no massive population movements associated with the spread of Hottentot pastoralism throughout southern Africa.     

Clumping and dispersal of chloroplasts in succulent plants

Plants have evolved various photoprotective mechanisms to mitigate photodamage. Here we report the diurnal movement of chloroplasts in the leaves of succulent crassulacean acid metabolism (CAM) plants under combined light and water stress. In leaves of water-stressed plants, the chloroplasts became densely clumped in one or sometimes two areas in the cytoplasm under light and dispersed during darkness. The chloroplast clumping resulted in leaf optical changes, with a decrease in absorptance and an increase in transmittance. The plant stress hormone abscisic acid induced chloroplast clumping in the leaf cells under light. We suggest that the marked chloroplast movement in these CAM plants is a photoprotective strategy used by the plants subjected to severe water stress.Abbreviations ABA Abscisic acid - CAM Crassulacean acid metabolism     

The Southern African orchid flora: composition, sources and endemism

Aim The Southern African orchid flora is taxonomically well known, but the biogeographical and diversity patterns have not yet been analysed. In particular, we want to establish whether (a) it is, like the Southern African flora in general, more diverse than would be expected from its latitude and area; (b) it is an African flora, or whether it contains palaeoendemic relicts of a Gondwanan orchid flora; (c) the diversity and endemism in the orchid flora is concentrated in particular biomes and habitat types; and (d) the patterns of endemism in the flora can be accounted for by current environmental parameters, or whether we need to invoke historical explanations. Location Southern Africa. Methods We used the recent floristic account of the Southern African orchids, in conjunction with a data base of over 14,642 herbarium records, to assign the species and subspecies of Southern African orchids to biomes, habitats, and clades. We explored the relationship between the number and endemism of entities (species, subspecies and varieties) and the biomes and habitats. We compared the richness of this flora with that of 31 other regions from all continents and latitudes, to establish whether the Southern African orchid flora is richer or poorer than expected. We assigned the Southern African orchid species to 16 monophyletic clades and mapped the global distribution of these clades to establish the continental affinities of the flora. Main conclusions The Southern African orchid flora is not any more diverse than could be expected from its latitude or area, while the two tropical African floras included were less diverse than expected. Latitude is an excellent predictor of regional orchid species richness; this might indicate that available habitat is more important for orchid diversity than gross area available, since latitude is probably correlated with the extent of suitable habitat. The Southern African orchid flora is clearly an African flora, since all clades are also found in tropical Africa, while many of them are absent from the Americas or Asia. Conversely, while most African orchid clades are also found in Southern Africa, both the Americas and Asia contain many clades absent from Africa. The distribution of orchid entities among the biomes in Southern Africa is very uneven, with two of the seven biomes totally devoid of orchids. Habitats and biomes that have no equivalent in tropical Africa are high in endemism, and habitats and biomes which are also well developed in tropical Africa are low in endemism. Endemism appears largely explained in terms of modern habitats. However, two patterns (the high endemism in the Succulent Karoo and the lack of endemism in the southern Cape among epiphytic orchids) may also be explained in terms of Quaternary climatic changes.     

Developing wetland inventories in southern Africa: A review

The status of wetland inventory effort and availability of maps and other data sources is reviewed for the ten countries of southern Africa: Angola, Bostwana, Lesotho, Malawi, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe. The aims and strategies for inventory are discussed and the main survey methods compared. Prior to commissioning new inventory work careful collation of existing maps and imagery is recommended together with targeting of strategic inventory at Province level, reserving high resolution effort only for certain important sites.     

Host range and distribution of small mammal fleas in South Africa,with a focus on species of medical and veterinary importance

The host range and distribution of flea species on rodents and insectivores across multiple vegetation types in South Africa were investigated. Habitat suitability for flea species considered as important vectors of disease in humans and domestic animals was modelled. Data originated from fleas that were recovered from small mammals captured at 29 localities during 2009–2013 and published literature searched for flea records. Climate‐based predictor variables, widely used in arthropod vector distribution, were selected and habitat suitability modelled for 10 flea vector species. A total of 2469 flea individuals representing 33 species and subspecies were collected from 1185 small mammals. Ten of each of the flea and rodent species are plague vectors and reservoirs, respectively. Multiple novel flea–host associations and locality records were noted. Three vector species were recorded from insectivores. Geographic distributions of flea species ranged from broad, across‐biome distributions to narrower distributions within one or two biomes. Habitat suitability models performed excellently for the majority of flea vectors and identified regions of summer and all‐year rainfall as representing suitable habitats for most vector species. Current knowledge of vector and disease ecology can benefit from similar sampling approaches that will be important not only for South Africa, but also for the sub‐region.     

The distribution and biology of nomadic birds in the Karoo, South Africa

Dryland nomadic bird species, as a proportion of all bird species in a biome in southern Africa, are highest in the arid grassland and arid and semi-arid Karoo in South Africa. Nomadic birds, of which the most widespread species is the greybacked finchlark Eremopterix verticalis (Smith), are most frequently observed in the north-central and north western Nama Karoo. The species richness of nomadic species is inversely correlated with species richness of all bird species in the Karoo. Since the distribution of nomadic birds is in areas where rainfall is patchy, low (<250 mm per year) and aseasonal, this supports the idea that fewer species are able to cope with resources that are patchy in time and space, and that there has been selection for nomadism in the species that are able to use patchy environments. Species richness and abundance of nomadic birds is negatively correlated with rainfall amount but positively correlated with the coefficient of variation of the rainfall and with rainfall in autumn. The frequency of nomadic birds is inversely correlated with altitude range; nomadic species are most often recorded in structurally simple habitats (shrubland and grassland) on open plains. Most nomadic bird species in the Karoo are granivorous. Perennial desert grasses are important components of the habitat and diet of small nomadic granivores, and also provide nest sites and nest material. Nomadic birds can breed throughout the year, without a clearly defined ‘season’ in both the Succulent and Nama Karoo. Average clutch sizes do not differ significantly between resident and all nomadic species in the arid and semi-arid Karoo. Nomadism is an evolutionary stable strategy for individual species only when extremes in environmental conditions are frequent enough, and unpredictable enough, to maintain movements to high resource patches or to maintain dispersal away from low resource patches. If high rainfall years are too regular or infrequent, or peaks in fluctuations of resources in the environment too low, or rainfall patches are randomly distributed, nomadism would not be maintained as part of the individual behaviour pattern.     

Taxonomy and distribution of some katydids (Orthoptera Tettigoniidae) from tropical Africa

Results of the study of specimens collected in tropical Africa and preserved in different European collections and museums are reported and extensively illustrated. The following three new species are described: Horatosphaga aethiopica sp. n., Dapanera occulta sp. n. and Cestromoecha laeglae sp. n. In addition, new diagnostic characters or distributional data for Ruspolia differens (Serville, 1838), Thyridorhoptrum senegalense Krauss, 1877, Horatosphaga leggei (Kirby, 1909), Horatosphaga linearis (Rehn, 1910), Preussia lobatipes Karsch, 1890 and Dapanera eidmanni Ebner, 1943 are reported. Finally, Symmetropleura plana (Walker, 1869) is proposed to be transferred to the genus Symmetrokarschia Massa, 2015, Conocephalus carbonarius (Redtenbacher, 1891) to the genus Thyridorhoptrum Rehn & Hebard, 1915; the genus Gonatoxia Karsch, 1889 is proposed to be synonymized with Dapanera Karsch, 1889.     

The radiation of the Cape flora, southern Africa

The flora of the south-western tip of southern Africa, the Cape flora, with some 9000 species in an area of 90,000 km2 is much more speciose than can be expected from its area or latitude, and is comparable to that expected from the most diverse equatorial areas. The endemism of almost 70%, on the other hand, is comparable to that found on islands. This high endemism is accounted for by the ecological and geographical isolation of the Cape Floristic Region, but explanations for the high species richness are not so easily found. The high species richness is accentuated when its taxonomic distribution is investigated: almost half of the total species richness of the area is accounted for by 33 'Cape floral clades'. These are clades which may have initially diversified in the region, and of which at least half the species are still found in the Cape Floristic Region. Such a high contribution by a very small number of clades is typical of island floras, not of mainland floras. The start of the radiation of these clades has been dated by molecular clock techniques to between 18 million years ago (Mya) (Pelargonium) and 8 Mya (Phylica), but only six radiations have been dated to date. The fossil evidence for the dating of the radiation is shown to be largely speculative. The Cenozoic environmental history of southern Africa is reviewed in search of possible triggers for the radiations, climatic changes emerge as the most likely candidate. Due to a very poor fossil record, the climatic history has to be inferred from larger scale patterns, these suggest large-scale fluctuations between summer wet (Palaeocene, Early Miocene) and summer dry climates (Oligocene, Middle Miocene to present). The massive speciation in the Cape flora might be accounted for by the diverse limitations to gene flow (dissected landscapes, pollinator specialisation, long flowering times allowing much phenological specialisation), as well as a richly complex environment providing a diversity of selective forces (geographically variable climate, much altitude variation, different soil types, rocky terrain providing many micro-niches, and regular fires providing both intermediate disturbances, as well as different ways of surviving the fires). However, much of this is based on correlation, and there is a great need for (a) experimental testing of the proposed speciation mechanisms, (b) more molecular clock estimates of the age and pattern of the radiations, and (c) more fossil evidence bearing on the past climates.