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.     

Community structure on unusual habitat islands: quartz-fields in the Succulent Karoo,South Africa

Quartz fields are edaphically arid, azonal habitats occurring under different macroclimatic conditions in several arid regions of southern Africa. They are the exclusive home of 142 plant species of which ca. 70% are local or regional endemics. This paper is an analysis of the quartz-field floras and growth form-soil relationships in two quartz field regions: the Knersvlakte in the Namaqualand-Namib Domain of the Succulent Karoo, and the western Little Karoo in the Southern Karoo Domain. The Knersvlakte supported 52 quartz-field specialists of which 39 were endemic to the region. Corresponding data for the Little Karoo were 11, and 10 species, respectively. In both regions, the average canopy cover on the quartz-field relevés was ca. 8%, and more than half of this comprised contracted, succulent nanochamaephytes. Cover and vegetation stature were markedly higher on adjacent zonal habitats. Quartz fields in both regions supported a similar array of compact, subglobose and subterranean nanochamaephytes, as evidenced by convergent patterns in two distantly related genera (Argyroderma N. E. Brown and Gibbaeum (Haworth) N. E. Brown, both Mesembryanthemaceae), endemic to the Knersvlakte and largely restricted to the Little Karoo, respectively. Analyses of vegetational and edaphic data of quartz fields and adjacent, zonal habitats were carried out using multivariate direct gradient analysis (Canonical Correspondence Analysis) in order to identify those factors that control the peculiar composition of growth forms on quartz fields. The results revealed highly similar patterns of growth form composition in relation to similar edaphic gradients in both regions. In general, the soils of quartz fields were shallower compared to those of adjacent zonal habitats. In both regions, two different groups of quartz-field edaphic habitats, representing extremes of a continuum, were identified. Group 1 was characterized by high salt content, neutral to slightly acid soil pH, and low stone content. Group 2 was characterized by low salt content, low soil pH, and high stone content. Group 1 quartz fields are the most edaphically arid habitats and support the highest relative cover and diversity of subglobose and subterranean chamaephytes. The combination of reduced competition from larger growth forms, shallow soils and high soil salinity, represents a regionally unusual selective regime. Some succulent lineages in the Mesembryanthemaceae have undergone diversification which has resulted in the fine-scale discrimination of subtle edaphic gradients within the saline quartz-patch habitats. Reliable seasonal rainfall and reduced thermal stress have also played a role in the evolution of quartz patch specialists.     

Effect of grazing on vegetation and soil of the heuweltjieveld in the Succulent Karoo,South Africa

We asked how historical and recent grazing intensity affect the patchy landscape of the heuweltjieveld in the semi-arid biodiversity hotspot Succulent Karoo. The study was carried out on a communal farmland 80 km south-west of Springbok, in Namaqualand. Heuweltjies are roughly circular earth mounds that are regularly distributed in this landscape. We sampled plant species and life-form composition, diversity measures, habitat and soil variables in 100 m² plots, placed in three visually distinguishable heuweltjie zones (centre, fringe, and matrix) and distributed across grazing camps with different recent and historic grazing intensities. Differences between heuweltjie zones were assessed with ANOVAs and multiple linear regressions. The effect of past and recent grazing intensity on soil and plant variables was analysed by Generalized Linear Models for each heuweltjie zone separately. The three zones constituted clearly distinguishable units in terms of vegetation and soil characteristics. Soil pH and cover of annual plants increased from matrix to centres, while total vegetation cover, species richness and perennial plant cover decreased in the same direction. Historic (pre-2000) grazing patterns had the strongest effects on fringes, showing the strongest soil and vegetation-related signs of overutilization with increased stocking density. Centres showed signs of overutilization irrespective of the stocking density. The much shorter exposure to recent grazing pattern (post-2000), which was nearly inverse to the historic grazing pattern, showed increase of vegetation cover (centres) and species richness (matrix) with recent grazing intensity. We interpret these effects as still visible responses of the lower grazing intensity in these camps during the historic period. No recovery under recent grazing was observed at any of the zones. We conclude that irrespective of their conducive growing conditions, once transformed to a disturbed state, heuweltjie centres recover slowly, whereas the less impacted soil and vegetation of fringes are more responsive than centres and matrix.     

Effects of Multispecies Clumping on Survival of Three Succulent Plant Species Translocated onto Mine Spoil in the Succulent Karoo Desert, South Africa

Salvaging from premining areas and translocation of succulent plants has been investigated for the revegetation of gypsiferous mine spoil in an arid region of South Africa. Given that facilitation effects are thought to outweigh competition effects in harsh environments, we hypothesized that the survival of translocated succulents would be higher when planted in multispecies clumps than alone and that the growth rate (measured as stem extension) and fruit‐set would be greater for plants in clumps than for those planted alone. Two leaf‐succulent species (Aridaria noctiflora ssp. noctiflora and Drosanthemum deciduum) and one stem‐succulent species (Psilocaulon dinteri) were salvaged from the area destined for mining and translocated onto the mine spoil. These plants were planted either in a multispecies clump of the three species together or alone. One year after translocation, 67% of the plants survived. It was also found that the succulents used in these experiments survived in higher numbers when planted alone. Due to the similar root morphology of D. deciduum and P. dinteri, they competed for resources instead of facilitating each other's establishment. The results were variable for each of the species used, and neither growth nor seed‐set was improved by clumping. These findings would also possibly vary from year to year with different abiotic conditions.     

The Roggeveldberge — Notes on a botanically hot area on a cold corner of the southern Great Escarpment, South Africa

Despite the Roggeveldberge comprising the montane heart of the celebrated Hantam-Roggeveld Centre of Plant Endemism (HRC), this section of the southern Great Escarpment in South Africa is botanically poorly known. A detailed physical, historical and phytogeographical overview of the Roggeveldberge is thus presented, and a checklist of 513 plant taxa is provided as a contribution towards a more complete flora for the Roggeveld-Komsberg Escarpment. The HRC is considered in detail, including various delimitations, and all species purported to be endemic to the HRC are reviewed and separated into actual endemics, near-endemics and species incorrectly considered as endemic.     

Floristics of the Dunbar Valley serpentinite site, Songimvelo Game Reserve, South Africa

The Dunbar Valley serpentinite outcrop has a flora comprising 254 taxa (species and below) in 172 genera and 63 families. Dunbar Valley has more species than other studied serpentinite sites in the Barberton Greenstone Belt (BGB). The genus Senecio is the most speciose genus in the BGB. The level of species endemism at Dunbar Valley is 2.0%. Most of the serpentinite endemics in the BGB show phytogeographical affinities with the Sudano-Zambezian Region. Six modified-Whittaker plots, three on serpentinite and three on non-serpentinite soils, were sampled. Sørenson's index was 0.312, indicating low similarity in species between serpentinite and non-serpentinite sites (β-diversity) at Dunbar Valley, as at other sites in the BGB. α-diversity (using the Shannon–Wiener index) for the serpentinite was 2.631 ± 0.901 and for the non-serpentinite, 2.886 ± 0.130. However, a Student's t -test showed no significant difference in α-diversity between the two habitats. There was also no significant difference in species richness between serpentinite and non-serpentinite sites. Total species showed negative correlations with total nickel, altitude and serpentinite outcrop size (area) for six sites in the BGB. Number of endemic taxa showed no correlation with environmental variables.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 271–285.     

Competitive and facilitative relationships among three shrub species,and the role of browsing intensity and rooting depth in the Succulent Karoo,South Africa

Abstract. The nearest‐neighbour technique is used to infer competition and facilitation between the three most abundant species in a semi‐arid region of western South Africa. Relationships among the shrubs Leipoldtia schultzei and Ruschia robusta, which are leaf‐succulent members of the Mesembryanthemaceae (‘mesembs’) and Hirpicium alienatum a non‐succulent Asteraceae, were compared on two adjacent sites with different histories of browsing intensity. Competition was more prevalent and more important than facilitation. The only evidence for facilitation was found at the heavily‐browsed site where the palatable Hirpicium was larger under the unpalatable Leipoldtia. Generally the prevalence and importance of competition was reduced at the heavily‐browsed site. Strong evidence was obtained for intraspecific competition in each of the three species; also, competition was evident between the two mesembs, where Leipoldtia was competitively dominant over Ruschia, although neither species inhibited Hirpicium. Minimal competition between the mesembs and the asteraceous shrub was interpreted in terms of differentiation in rooting depth, and competition within the mesembs, in terms of overlap in rooting depth. The mesembs had the bulk of their roots in the top 5 cm of soil, while the asteraceous shrub had the bulk of its roots, and all its fine roots, at greater depths. The shallow‐rooted morphology of the mesembs is well adapted to utilize small rainfall events, which occur frequently in the Succulent Karoo, and do not penetrate the soil deeply. Modifications of existing methods are applied for analysing nearest‐neighbour interactions.     

Insights into the biodiversity of the Succulent Karoo hotspot of South Africa: the population genetics of a rare and endemic halictid bee,Patellapis doleritica

Population genetic analyses are especially relevant for species considered threatened or highly endemic and for which other forms of biological information are lacking. Patellapis doleritica is a recently described communally nesting halictid bee of conservation concern because it is rare and endemic to the Succulent Karoo of South Africa. Moreover, its dispersal is considered to be restricted by its specialised nesting requirements and inclement weather conditions during its limited annual flight period, traits which may be common to other bee species of the region. We hypothesised that gene flow in P. doleritica was low, leading to marked genetic differentiation. Using 7 microsatellites, we investigated its mating and population genetic structure in 258 individuals (171 females and 87 males) from 7 populations spanning most of its known range. Deviation from Hardy–Weinberg equilibrium (F_IS = + 0.254) suggested P. doleritica to be inbred, as in many other communal nesting bee species. Global F_ST (0.028) and global G′_ST (0.216) revealed modest but significant differentiation between most populations, even across the very limited range of the species (ca. 25 km), with one genetically extreme outlier population. Despite inbreeding, we detected a surprisingly low frequency of diploid males (2 %). Patellapis doleritica nevertheless deserves special conservation attention since it is an endemic species with a low overall abundance and therefore possibly prone to environmental change and local extinction.     

The importance of monkey beetle (Scarabaeidae: Hopliini) pollination for Aizoaceae and Asteraceae in grazed and ungrazed areas at Paulshoek, Succulent Karoo, South Africa

The relative importance of monkey beetles (Hopliini, Scarabeidae) as pollinators of Asteraceae and Aizoaceae in the Succulent Karoo as well as the influence of livestock grazing on their abundance and diversity was investigated. Hopliine beetles proved to be the, or among the, most abundant flower visitors of 12 investigated plant species. However, during single flower observations at three Aizoaceae species, bees (Apoidea), bee flies (Bombyliidae) and pollen wasps (Masaridae) were the most frequent flower visitors. However, monkey beetles carried the highest Asteraceae and Aizoaceae pollen loads, and are therefore considered to play a vital role in the pollination of these two families. Abundance, species richness and diversity of Hopliini did not appear to be heavily affected by livestock grazing. Annual variation in the composition of monkey beetle populations was more dramatic. Still, some species showed higher abundances on heavily grazed rangeland while others only occurred under low grazing pressure. It is presumed that changes in the composition of the vegetation, especially the observed decrease of perennial plants in favour of annuals and geophytes (Todd and Hoffman 1999) could in turn affect the composition of monkey beetle assemblages.     

神农架北坡堵河源自然保护区植物多样性研究

为建立自然保护区,保护神农架北坡堵河源地区丰富的生物多样性资源,并为神农架地区植物起源、植被演替以及生物多样性和濒危物种状况深入、准确评价提供基础数据,2002年,采用路线调查与标准地调查相结合的方法,结合历史资料,对堵河源地区的野生植物资源进行了实地野外调查研究,分析了植物区系和植物多样性及其受威胁状况。结果表明:(1)该保护区具有丰富的植物物种多样性,有维管束植物174科730属1733种(含亚种、变种),其中,国家重点保护野生植物有23种;(2)该区植物区系成分具有复杂性、多样性,拥有中国种子植物分布区类型属级水平的全部类型,但明显偏重于温带性质。且植物区系起源古老,有中国特有分布属37个,其中,古特有属27个;(3)该区具有丰富的植被类型,包括4个植被型组,8个植被型,26个群系,具有较高的保护和科研价值。同时针对保护区植物多样性受威胁状况,提出了植被恢复、加强防火设备设施建设、生态移民、保护珍稀濒危植物及社区共管是保护植物多样性的有效措施。     

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.     

Species turnover: the case of stream amphibians of rainforests in the Western Ghats, southern India

We examined species turnover in stream amphibians in rainforest in two hill ranges (Ashambu and Anamalai Hills) in the Western Ghats in south India. In each hill range, six stream segments (100 m in length) belonging to three drainage or rivers were surveyed three to four times in three seasons over 1 year. Species turnover (using 1-Sorenson's index) was estimated between all possible pairs of sites at three spatial scales – within drainage, between drainage and between hill ranges. Similar matrices were also developed for altitudinal difference and geographic distance between sites. A total of 30 species in four families were recorded from 3681 individuals. The hill ranges differed significantly in the composition of the stream community at both the species and family levels. Within the hill range, species turnover was correlated with altitudinal difference and not with geographic distance. Anamalai Hills had a greater species turnover than Ashambu Hills, both within and between drainage. There was also a high turnover between these two hill ranges, with only two shared species. This turnover explains the fact that only 30–40 species have been reported from different hill ranges, although regional diversity is high with about 130 species. The turnover also predicts that several undetected species should occur in hill ranges and drainage that have not been surveyed. The conservation model for mammals and birds, consisting of a few large protected areas, may not adequately address the conservation requirements of amphibians. Protection of rainforest frogs may require many protected areas in different drainages.     

Density‐dependent regulation of the critically endangered black rhinoceros population in Ithala Game Reserve,South Africa

Ensuring the persistence of populations of endangered species requires an understanding of, and response to, the causes of population declines. Species occurring in small populations are vulnerable to stochastic problems that are environmental, demographic, or genetic in nature and can reduce survival as much as the deterministic threats of habitat degradation. Critically endangered black rhinoceros (Diceros bicornis) populations declined throughout Africa since 1960, with numbers steadily increasing at a continental level, but remaining lower than three generations ago. However, size, demographics, trends, and factors affecting these, are poorly known. We used 18 years (1990–2008) of long‐term sightings data from Ithala Game Reserve, KwaZulu‐Natal, South Africa, to determine population estimates, growth rate and fecundity over time, as well as sex and age structure and age‐specific probabilities of survival. Calf survivorship between the ages of 0 and 1 year was 74% for females and 94% for males. Age‐specific survivorship for both sexes was significantly higher from yearling to subadult age‐classes (1–6 years) than for adults (7–30 years). The most frequent cause of mortality was attributed to unknown causes while fighting injuries was recorded as the second most common cause of mortality, particularly among subadult and adult males. There was no significant difference in the sex ratios at birth, although the proportion of females in the population was 0.58. There was strong evidence for density‐dependent regulation, with density in conception year a key driver of population performance (birth rate). The population does not appear to be at ecological carrying capacity; however, social effects are delaying conception. To mitigate density‐dependent social effects, we recommend an adaptive management strategy of pre‐selecting individuals for removal from the reserve, so as to maintain stability in the social organization of the population.     

Species turnover of monkey beetles (Scarabaeidae: Hopliini) along environmental and disturbance gradients in the Namaqualand region of the succulent Karoo, South Africa

Species turnover of monkey beetle (Scarabaeidae: Hopliini) assemblages along disturbance and environmental gradients was examined at three sites within the arid, winter rainfall Namaqualand region of the succulent Karoo, South Africa. At each site two study plots with comparable vegetation and soils but contrasting management (grazing) histories were chosen, the disturbed sites having fewer perennial shrubs and generally more annuals and bare ground. Beetles collected using coloured pan-traps showed a consistently higher abundance in disturbed sites. Lepithrix, Denticnema and Heterochelus had higher numbers in disturbed plots, while Peritrichia numbers were lower in disturbed areas. Measures of species richness and diversity were consistently higher in the undisturbed sites. Distinctive assemblages of monkey beetles and plants occurred at each site. A high compositional turnover ( diversity) was recorded for both monkey beetles and plants along a rainfall gradient; between-site diversity values ranged from 0.7 to 0.8 (out of a maximum of 1.0). Species turnover of beetles was higher between the disturbed sites along the environmental gradient than the corresponding undisturbed sites. The high monkey beetle species turnover is probably linked to the high plant species turnover, a distinctive feature of succulent Karoo landscapes. Monkey beetles are useful indicators of overgrazing disturbance in Namaqualand, as their pollinator guilds are apparently disrupted by overgrazing. A shift away from perennial and bulb pollinator guilds towards those favouring weedy annuals was observed in disturbed areas. The consequences to ecosystem processes due to the effects of disturbance on monkey beetle communities and the role of monkey beetles as indicators of disturbance is discussed, as well as the implications of disturbance on monkey beetle pollination guilds.     

A new species of Polygala (Polygalaceae) from ultramafic soils in Sekhukhuneland,South Africa,with notes on its ecology

Polygala sekhukhuniensis Retief, Siebert & A.E.Van Wyk (Polygala; section Polygala; subsection Heterolophus), a new species with a restricted range in Sekhukhuneland, South Africa, is described, illustrated and compared with other members of the genus. It is a dwarf shrub that can be distinguished by its much-branched habit, sparsely flowered inflorescences, pink alae with darker pink veins, brown to black seed testa, and oblate pollen grains with pronounced opercula. Geographically, P. sekhukhuniensis is confined to heavily eroded localized sites, a natural geomorphological feature of some of the highly water-dispersible soils derived from ultramafic rocks in the valleys of the Steelpoort River and its tributaries in the Sekhukhuneland Centre of Plant Endemism. P. sekhukhuniensis is a calciotrophic excluder of heavy metals that accumulates Ca in its leaves. It is ecologically compared with co-occurring species of Polygala on ultramafic-derived soil.     

Natural plant species inventory of hotspot areas in Arabian Peninsula: Southwest Al-Baha region,Saudi Arabia

Plant hotspot areas are the areas that are very rich in plant species diversity. These areas have a priority for conservation. To highlight the plant diversity for nature conservation purposes a case study in Al Baha region, Saudi Arabia is presented, in which the importance of the natural vegetation and flora of one of the hotspot areas of Saudi Arabia is evaluated through the explanation of its natural plant species. A survey study has been conducted in an area of 167.6 km2, a 97 sample each with 20X20 m were laid out covering the whole ecological zones of the study site. Data of flora, vegetation cover and topography were gathered from each sample site. The study revealed about 319 plant species belonging to 228 genera and 75 families. Two species were found endemic to Saudi Arabia, 14 were endemics to Arabian Peninsula, and five were regional endemics that are only found in East Africa and Arabian Peninsula, while 39 species are rare and endangered.     

珠穆朗玛峰国家级自然保护区密叶红豆杉所在群落的特征及区系分析

基于野外15个群落样方物种调查,对珠穆朗玛峰国家级自然保护区密叶红豆杉(Taxus contorta Griffith)所在群落的结构、物种多样性、区系地理成分及密叶红豆杉的年龄结构进行研究。结果显示:(1)调查样地内共有维管植物53科116属154种,以蔷薇科、毛茛科、菊科、百合科等占优势;在科、属级区系分析中,热带成分分别占46.2%、18.0%,温带成分分别占53.8%、82.0%。(2)密叶红豆杉所在群落的建群种分别为乔松(Pinus wallichiana A.B.Jackson)、云南铁杉(Tsuga dumosa(D.Don)Eichler)和喜马拉雅冷杉(Abies spectabilis(D.Don)Spach),密叶红豆杉为下层乔木的重要伴生成分或优势成分。(3)从各群落内部比较来看,草本层的Margalef指数和Simpson指数在3个群落中都最高,说明群落的草本层物种多样性最丰富,数量也最多;从群落间比较来看,乔松林灌木层和草本层的Margalef指数最高,乔木层的最低。(4)在3种群落中密叶红豆杉数量都呈增长趋势,且幼苗库较充足,在云南铁杉林和喜马拉雅冷杉林中,密叶红豆杉从幼苗到幼树的生长过程经历了较强的环境筛选。     

湖北星斗山国家级自然保护区种子植物区系研究

于2013-2015年对湖北星斗山国家级自然保护区进行多次全面野外调查,采集植物标本3441号,并对其种子植物区系特点进行了研究。结果显示,该保护区共有野生种子植物138科678属1617种,其中裸子植物6科15属19种,被子植物132科663属1598种。区系分析表明,热带科(66.06%)多于温带科(33.94%),这与该区曾经历过古热带的历史背景相吻合;属级水平上则为温带属性(58.45%)强于热带(41.55%);中国特有种623个(占总种数的38.53%),均与四周区域所共有,反映出东西南北植物区系在此交汇渗透的态势;这些均表明该区种子植物区系成分不但复杂多样,而且还具有过渡区域的特点;保护区内植物区系还具有古老、残遗的特性,一些古老的残遗植物在该地有良好的原生群落分布;保护区内特有树种和珍稀濒危植物也很丰富,具有很大的保护价值。