Desertification,no‐change or alternative states: Can we trust simple models on livestock impact in dry rangelands?

Abstract. It is remarkable that after many thousands of years of pastoralism in arid and semi‐arid savannas, there is still no clear answer to the basic question: Does livestock grazing have more than a marginal effect on the dynamics of arid and semi‐arid land vegetation? A small study of semi‐arid savanna vegetation along a spatial gradient in grazing pressure, repeated three times over 19 years is used as a basis for discussing the behaviour of dry land vegetation under heavy grazing. Three basic theories are compared: (1) the theory that heavy grazing causes desertification; (2) the theory that heavy grazing causes no directed change in the vegetation of arid and semi‐arid rangelands; and (3) the theory that heavy grazing leads to a switch between alternative states of vegetation. On the basis of the current data the first two theories are rejected, but the conclusion is that there is much evidence for a more complex behaviour of dry rangelands under grazing than what is accommodated in any of the three theories. Probably, site‐specific properties including interactive and indirect effects of herbivory in the system are important for vegetation development under grazing in arid and semi‐arid rangelands.     

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.     

Stable isotopes reveal regional movement patterns in an endangered bustard

Stable isotope analysis is a valuable technique to infer animal movement between isotopically distinct landscapes. For birds in terrestrial systems, it is usually only applied at continental scales, often relying on global isotopic patterns. In contrast, we used this technique to investigate movement patterns of Ludwig's bustard (Neotis ludwigii) at a regional scale, where such information is needed to improve the conservation status of this species. We analysed carbon and nitrogen isotopic compositions of feathers from bustards across two biomes of the semi‐arid rangelands of the Karoo, South Africa, to investigate movement and explore sex and age movement strategy differences. We used a linear discriminant function analysis based on growing feathers to classify fully grown feathers to a Succulent or Nama Karoo biome origin. Six of 12 birds for which all primary feathers were analysed had at least one feather classified as having grown in the Succulent Karoo, supporting the theory that these birds are partial migrants. Feathers from two satellite‐tracked bustards broadly supported the conclusions of the analysis, although food base differences resulting from local rainfall variation probably obscured geographic signals at finer scales. There was no apparent difference in movement strategies between the sexes, but juvenile feathers were almost exclusively assigned to the Nama Karoo, suggesting that most breeding occurs in this biome. Adult and juvenile feathers also had significantly different isotope ratios, which could relate to diet or to differing metabolic processes. This study demonstrates that with a good understanding of the system, carbon and nitrogen stable isotopes can be useful to infer general movement patterns of birds at a regional level.     

Boom means bust: interactions between the El Niño/Southern Oscillation (ENSO), rainfall and the processes threatening mammal species in arid Australia

We collated an environmental history for a 8580 km² study area in the Simpson Desert, Australia. Quantitative and qualitative data on climate, land-use, fire history and ecosystem dynamics were used to construct a chronology of processes threatening terrestrial mammal species. Over the last 150 years there has been the transition in land tenure from a hunter–gatherer economy to pastoralism, the loss of 11 mammal species, the cessation of small scale burning by Aboriginal people and the introduction of the fox and cat. Annual rainfall was highly variable and was influenced by the phase of the El Niño Southern Oscillation (ENSO). Irruptions of rodents, marked increases in the populations of native and introduced predators and extensive wildfires were associated with the La Niña phase of ENSO and occurred when rain-year (July–June) rainfall approached or exceeded the 90th percentile of the historical rainfall distribution. Large rainfall events in arid Australia have been viewed traditionally as the ‘boom’ times that benefit wildlife and pastoral production. However, because of hyper-predation and the risk of wildfire, we show that the years including and immediately following flooding rains should be identified as critical, or ‘bust’ periods for wildlife and conservation management. ENSO related climatic forecasts appear to be useful cues which can be incorporated into fire and predator management strategies in arid Australia. Studies such as this, which utilise a broad range of data types across extensive areas, can identify the timing and potential of threatening process not possible using contemporary studies alone.     

Alien plant assemblages near roads in arid and semi-arid South Africa

Herbaceous and woody alien plants visible from a moving vehicle were recorded along 1 km roadside transects at 5 km intervals over a distance of 5869 km in the semi-arid and arid Fynbos, Succulent Karoo, Nama Karoo and Arid Savanna (Kalahari) biomes in South Africa. Each 1 km transect was classified by biome and vegetation type, mean annual rainfall, rainfall seasonality, soil surface type and landuse adjoining the roadside. Although travelling speed affected the range and frequency of plant species observed, the method was repeatable at a speed of 100 km h^?1. Alien plants occurred in 98% of 119 Fynbos, 81% of 204 Succulent Karoo, 72% of 661 Nama Karoo, 47% of 171 Arid Savanna and 100% of seventeen Grassland transects. Alien species richness per site was correlated with mean annual rainfall, but in all regions, sites adjacent to cultivation had significantly more alien species than sites adjoining rangeland. The alien plant assemblage of the arid winter-rainfall Succulent Karoo included species from mesic winter-rainfall lowland Fynbos and the arid Nama Karoo receives unseasonal rainfall. The frequencies of Prosopis spp., Atriplex spp. and Opuntia ficus-indica were not significantly greater near cultivation, and these perennial plants, all of which are dispersed by indigenous and domestic animals, can invade natural rangeland in arid and semi-arid southern Africa.     

How the west was once: vegetation change in south-west Queensland from 1930 to 1995

Aim Conflicting perceptions of past and present rangeland condition and limited historical data have led to debate regarding the management of vegetation in pastoral landscapes both internationally and in Australia. In light of this controversy we have sought to provide empirical evidence to determine the trajectory of vegetational change in a semi‐arid rangeland for a significant portion of the 20th century using a suite of proxy measures. Location Ambathala Station, approximately 780 km west of Brisbane, in the semi‐arid rangelands of south‐western Queensland, Australia. Methods We excavated stratified deposits of sheep manure which had accumulated beneath a shearing shed between the years 1930 and 1995. Multi‐proxy data, including pollen and leaf cuticle analyses and analysis of historical aerial photography were coupled with a fine resolution radiocarbon chronology to generate a near annual history of vegetation on the property and local area. Results Aerial photography indicates that minor (< 5%) increases in the density of woody vegetation took place between 1951 and 1994 in two thirds of the study area not subjected to clearing. Areas that were selectively or entirely cleared prior to the 1950s (approximately 16% of the study area) had recovered to almost 60% of their original cover by the 1994 photo period. This slight thickening is only partially evident from pollen and leaf cuticle analyses of sheep faeces. Very little change in vegetation is revealed over the nearly 65 years based on the relative abundances of pollen taxonomic groups. Microhistological examination of sheep faeces provides evidence of dramatic changes in sheep diet. The majority of dietary changes are associated with climatic events of sustained above‐average rainfall or persistent drought. Most notable in the dietary analysis is the absence of grass during the first two decades of the record. Main conclusions In contrast to prevailing perceptions and limited research into long‐term vegetation change in the semi‐arid areas of eastern Australia, the record of vegetation change at the Ambathala shearing shed indicates only a minor increase in woody vegetation cover and no decrease in grass cover on the property over the 65 years of pastoral activity covered by the study. However, there are marked changes in the abundance of grass cuticles in sheep faeces. The appearance and persistence of grass in sheep diets from the late 1940s can be attributed to the effects of periods of high rainfall and possibly some clearing and thinning of vegetation. Lower stock numbers may have allowed grass to persist through later drought years. The relative abundances of major groups of plant pollen have not changed significantly over the past 65 years.     

An underground index of rangeland degradation: cicadas in arid southern Africa

Summary The prediction that density of long-lived, underground herbivores (Cicadidae) is a function of rangeland condition was examined in arid shrublands in the Karoo, South Africa. It was found that the density of adult cicadas was correlated with cover of deep-rooted, perennial plants. Differences in perennial plant cover were independent of soil type and rock cover. On degraded rangelands, where perennial plants had been replaced by ephemerals and short-lived plants, cicada densities were significantly depressed. We concluded that vegetation transformation by domestic livestock is likely to affect invertebrate populations and their vertebrate predators.     

Ecological correlates of small mammal community structure in the semi-arid Karoo, South Africa

The understanding of the determinants of small mammal community structure in arid and semiarid ecosystems is of importance, both in the light of the role that small mammals play, and the impact of livestock grazing on the flora of these systems. In a study aimed at identifying these determinants, small mammal assemblages and environmental features were quantified at six localities (a gradient of floristic structure, with constant annual rainfall) across the southern Karoo, South Africa. Stepwise variable regression indicated that small mammal diversity was correlated with plant and rock cover, as well as plant cover and horizontal foliage diversity at intermediate heights (40–60 cm). Initially, small mammal diversity increased with increasing plant cover, but decreased at cover levels greater than 30%. This relationship is similar to that found in other desert systems, although the peak in diversity found here is at higher levels of plant cover than found previously. I suggest that this higher peak may be owing to the lack of reliance on granivory by these animals, which are relatively omnivorous. This model may therefore explain the conflicting reports on the impact of livestock grazing on desert small mammals, with small mammal diversity decreasing with grazing below the peak, and increasing with grazing above the peak.     

Comparative patterns of phenology and growth form diversity in two winter rainfall deserts: the Succulent Karoo and Mojave Desert ecosystems

A comparative study of community structure and seasonal growth dynamics in the arid winter rainfall regions of the Succulent Karoo in South Africa and the Mojave Desert of the United States suggests that remarkably divergent patterns of resource use and resultant growth form diversity exist in regions with outwardly similar climatic regimes. An understanding of these divergent patterns in the two winter rainfall deserts allows predictions to be made on vegetation response to global change. Above-ground plant growth in the Succulent Karoo begins with the first significant rains in late summer and continues through winter because moderate minimum temperatures allow continued growth. These communities have low structural diversity above-ground, but also below-ground, where root systems commonly do not exceed 20 cm in depth. These shallow root systems harvest water from upper soil horizons soon after rain falls, and growth declines as rainfall decreases in late spring. In contrast, low temperatures during winter inhibit growth in the Mojave Desert until early spring at a time when a mean 74% of the hydrologic year precipitation (July-June) has already occurred. Thus species in this structurally diverse system rely on deeper stores of water for growth in spring and early summer. A global change scenario of a 2 to 4°C increase in mean annual temperature and increased summer rainfall in the Mojave desert would be expected to produce similar conditions in the Mojave Desert to those that exist in the Succulent Karoo today. Assuming no genetic constraints on phenotypic plasticity, this would suggest increased species diversity and a decline in structural diversity in the Mojave Desert over evolutionary time. Increased summer rainfall in the Succulent Karoo would be expected to lead to invasions of grasses and thus increased competitive pressure reducing community diversity.     

Population status and distribution modelling of the critically endangered riverine rabbit (Bunolagus monticularis)

The riverine rabbit (Bunolagus monticularis) is one of the most endangered mammals in the world due to fragmentation of its habitat in the semi‐arid Karoo region of South Africa, to which it is endemic. It is an umbrella conservation species for the riparian shrubland associated with the seasonal drainage system of the Karoo, where its presence is an indicator of ecosystem health. In this study, we analysed historical survey data to derive an improved assessment of the current B. monticularis population status and distribution. Geospatial analysis was conducted using geographical information systems, and distribution modelling was performed using Maxent. Extent of occurrence for the species is 54,227 km², and area of occupancy is 2943 km². Estimates of 157–207 mature individuals confirm an alarmingly small species population size, and it appears that no subpopulation has >50 mature individuals. Our findings thus support the continued classification of this species as ‘critically endangered’ under IUCN Red List criteria. However, with most remaining habitat occurring outside of protected areas, and with habitat loss being exacerbated by climate change, a viable conservation plan remains elusive.     

Edaphic and climatic history has driven current dung beetle species pool and assemblage structure across a transition zone in central South Africa

We investigate biogeographical, regional and sub‐regional‐scale responses of scarabaeine dung beetles to late Cenozoic changes in edaphic and climatic character that created a Savanna/Karoo transition zone in the Northern Cape, South Africa. Across a 50 200 km² study area, the Northern Cape species pool comprised six biogeographical groups defined from distribution across southern Africa. These species groups contributed in different proportions to five regional assemblages defined from structural differences across the transition zone. Towards transition zone peripheries, regional assemblage structure was more strongly correlated to sandiness dating from Miocene to Pliocene deposition (Kalahari), aridity dating from Pliocene to Pleistocene climatic change (Bushmanland Karoo), or cooler temperatures dating from Miocene to Pliocene uplift (Upper Karoo). Correlates of sub‐regional assemblages trended to intensification of dominant drivers towards regional peripheries. Drivers of central transition zone, regional assemblages (‘Gariep Karoo’, ‘Gariep Stony Karoo’) showed no dominance. Biogeographically, endemism dominates the Northern Cape transition zone: south‐west arid groups in Nama Karoo regions; Kalahari plus north‐east savanna groups in the Kalahari. Regionally, transition drives assemblage structure: unique variance, 60% in the Kalahari, 21–30% in four Nama Karoo regions; shared variance (overlap), 25–65% between Kalahari and warmer Karoo regions, 11–71% between mainly cooler Karoo regions.     

Multi-scale determinants of dung beetle assemblage structure across abiotic gradients of the Kalahari–Nama Karoo ecotone, South Africa

Aim Changing conditions across spatial gradients are primary determinants of biotic regions, local habitats, and distributional edges. We investigate how a climatic gradient and edaphic mosaic interact as multi‐scale drivers of spatial patterns in scarabaeine dung beetles. The patterns are tested for congruency with ecoregion and floral boundaries over the same gradient, as responses to physical factors often differ among higher taxa. Location Southern Africa and the Nama Karoo–Kalahari ecotone, Northern Cape, South Africa. Methods Data consisted of the climatic distributions of 104 species and their abundances at 223 sites in two ecoregions/floral biomes, four bioregions, and 13 vegetation units. Factor analyses determined the biogeographical composition of the species, and regional‐ to local‐scale patterns in species abundance structure. Hierarchical analysis of oblique factors determined the proportional contribution of spatial variance to patterns. One‐way anova was used to test for significant separation of patterns along factor axes. Stepwise multiple regression was used to determine correlations of five physical attributes with species richness, Shannon‐Wiener diversity, and factor loadings for the study sites. Results Four biogeographical influences overlap in the study region, although rank contribution declines from south‐west arid through north‐east savanna to widespread and south‐east highland taxa. Species abundance structure comprises five subregional patterns, two centred to the north‐east (Kalahari, Isolated Kalahari Dune) dominated by Kalahari influence, and three to the south‐west (Nama Karoo subdivisions: Bushmanland, ‘Upper’, ‘Stony Prieska’) dominated by south‐west arid influence. Kalahari deep sands are characterized especially by a warmer, moister climate, whereas the Nama Karoo mosaic of deep or stony soils is characterized especially by north‐west aridity (Bushmanland), south‐east cooler temperatures (‘Upper’), or excessively stony soils (‘Stony Prieska’). Four of the subregional patterns each comprised three localized patterns related primarily to relative stoniness, edge effects from geographical position, or incidence of rainfall. Species richness and diversity declined with decreasing rainfall and increasing stoniness. Main conclusions Climatic and edaphic factors are important multi‐scale determinants of spatial patterns in dung beetle assemblage structure, with edaphic factors becoming more important at local spatial scales. The patterns are roughly congruent with the Kalahari Savanna–Nama Karoo ecotone at the floral biome or ecoregion scale, but show limited coincidence with finer‐scale floral classification.     

Meteorological periodicities in zonal mean annual temperatures and precipitation totals

Abstract

A long‐period data set (1880–1970) of mean annual temperatures and precipitation totals for over 100 stations worldwide distributed was used to calculate time series of zonal means at 10 degree latitude bands. Power spectra of these zonally averaged series display the following features: over most of the earth zonal mean annual temperatures undergo long‐term climatic fluctuations (≥22 years).

Trends in rainfall totals are only detected in the zones 20°‐30°N and 40°‐50°N. As reported earlier for single stations, a quasi‐triennial oscillation (3–4 years) is seen also in zonal rainfall totals at most latitude belts. No 11‐years peak was observed both in temperature and precipitation spectra. A 22‐year peak is observed at equatorial zones and in the zone 40°‐50°N.     

干旱和半干旱地区灌木下土壤“肥岛”研究进展

世界许多干旱、半干旱地区草地生态系统逐渐被灌木生态系统取代,这种取代过程的发展是由灌木冠幅下土壤中“肥岛”的形成和灌木的扩散相互作用直接造成的,这种“肥岛”的形成和灌木的扩散之间的反馈作用能够改变该系统中的植被组成、结构和土壤养分分布格局,从而改变该生态系统的结构和功能,文中从“肥岛”的概念和形成机制出发,对“肥岛”现象形成的原因、“肥岛”现象的研究意义、研究方法、灌木扩散与“肥岛”之间的相互关系及生物对“肥岛”的响应进行了总结,并分析了“肥岛”研究中应注意的问题,希望为中国干旱、半干旱区域土地退化的成因和过程以及植被恢复的研究提供一些参考。     

Cenozoic environmental change in South America as indicated by mammalian body size distributions (cenograms)

A cenogram is a rank‐ordered body size distribution of non‐predatory terrestrial mammal species within a community. Studies of cenograms for modern faunas have shown that certain quantifiable attributes of cenograms are correlated with environmental variables such as rainfall and vegetation structure. Based on these correlations, cenograms of fossil communities have been used to infer palaeoenvironments and palaeoenvironmental variables. The present study uses cenogram statistics to interpret palaeoenvironmental conditions for eight Cenozoic South American mammal faunas, ranging from Eocene to Pleistocene in age. Body sizes for fossil taxa were taken either from the literature or were estimated using regressions of body size on molar length (or femoral bicondylar width) for modern mammals. Cenogram statistics are calculated for the eight fossil faunas and compared to similar statistics calculated for 16 modern South American mammal faunas, allowing palaeoenvironmental interpretations to be made. The palaeoenvironmental interpretations based on cenogram analyses sometimes support and sometimes contradict interpretations based on herbivore craniodental morphology (e.g. levels of hypsodonty). Simulations of expected errors in body size estimates for fossil taxa suggest that the discrepancies do not result primarily from erroneous body size estimates. It is possible that some of the incongruity in interpretations results from certain non‐analogue attributes of South American faunas during much of the Cenozoic (e.g. the relatively depauperate mammalian predator diversity prior to the Great American Biotic Interchange).     

Hierarchical processes define spatial pattern of avian assemblages restricted and endemic to the arid Karoo, South Africa

Abstract Aim To identify and quantify biotic and abiotic factors associated with the regional gradients in the distribution and abundance of bird communities restricted and endemic to the Succulent and Nama Karoo biomes of South Africa. Location The arid Nama and Succulent Karoo biomes in South Africa. Methods The quarter degree grid cell (QDGC) was used to extract environmental data, while the bird data previously atlased, was linked to the same geo‐referenced system, using a geographical information system (GIS). Bird species were grouped into different life‐history assemblages. A quantitative, systematic analysis of the different bird communities spanning the Karoo was undertaken to examine contributions of broad‐ and local‐scale physical environmental and biotic factors to regional variations in the species composition, using multivariate statistical and spatial analytical tools. These included two indirect gradient methods; principal components analysis (PCA) and detrended correspondence analysis (DCA), and two direct gradient methods; canonical correspondence analysis (CCA) and redundancy analysis (RDA). Results Principal components analysis results showed that the selected environmental variables accounted for about 85% of the variation in the region. The first two principal gradients defined regional temperature seasonality and variability especially in winter and summer. The third principal gradient mainly defined summer rainfall areas in association with the coefficient of variation of rain and regional primary production, while the fourth gradient defined winter rainfall areas, growth days and elements of landscape structure. CCA/RDA analysis produced shortened hierarchically ranked explanatory variables for each bird assemblage. Stepwise gradient analysis results showed summer rain, rainfall concentration, topographic heterogeneity and annual evapotranspiration, as the most important climate variables explaining species occurrence. Landscape, in terms of percentage transformation, morphology, coefficient of variation of primary productivity and distance between suitable habitat patches, were also important, but to a lesser degree. Total variation explained (TVE) by the supplied variables was between 23 and 37% of variation. Less than 20% of TVE was the intrinsic spatial component of environmental influence, indicating that any unmeasured factors were independent of spatial structuring. For all the eight bird assemblages, climate contributed most to TVE (24–57%). Landscape characteristics (human‐induced transformation, vegetation in terms of size if grassy clumps and the average distances between them) contributed theleast to TVE for all the different assemblages (0–6%), especially the granivorous assemblage where it was not significant at all (0%). Seasonal extremes and variability were more important in explaining species gradients than were annual climatic conditions, with the exception of annual potential evapotranspiration. Main conclusions This study was able to synthesize species environment relations at the broad scale and demonstrated the association of arid zone endemic species occurrence with climate extremes and seasonality. Given the predicted climate change scenarios for South Africa, this regional gradient study provides a quantitative ecological basis for finer scale modelling and analysis, developing regional strategies for conserving biodiversity as well as predicting and planning for the effects of global climate change. However, most importantly, it clearly showed that bird species restricted and endemic to the arid Karoo biome may be more sensitive to climate rather than vegetation structure as previously thought.     

Oscillations in large mammal populations: are they related to predation or rainfall?

Cyclic population dynamics is relatively common among populations of small mammals in high latitudes but is not yet established among African savanna ungulates. However, oscillations may be expected in large mammal populations subject to quasi‐periodic oscillations in regional rainfall. We evaluated evidence for environmentally entrained oscillations in a large‐mammal predator–prey system in Kruger National Park (KNP), South Africa, where rainfall exhibits quasi‐periodic oscillations. The evaluation is based on analysis of comparative changes in the abundance of twelve ungulate species throughout South Africa's KNP using population counts over the period 1965–1996. We present evidence suggesting that (i) twelve ungulate populations display cyclic variability with half‐periods ranging between 10 and 18 years, (ii) this variability was associated with lagged rainfall between 3 and 10 years back in the past for different ungulate species, and (iii) the ungulate species respond in contrasting ways to rainfall, with some reaching highest abundance during periods of low rainfall and others under conditions of high rainfall. These findings are not consistent with the response pattern we would expect if the population oscillations were driven directly by the rainfall influence on food availability. Instead they seem to be an outcome of predator–prey interactions, which are entrained by the effect of rainfall on habitat conditions affecting the relative susceptibility of the different ungulate species to predation.     

Are nurse‐protégé interactions more common among plants from arid environments?

Abstract. Harsh conditions in arid and semi‐arid environments make seedling establishment rare. Plant recruitment in arid environments often occurs only in years with above average rainfall or in safe sites under the canopy of nurse plants that provide shelter from high temperatures and low moisture. Associations of establishing seedlings with adult plants are referred to as nurse‐protégé interactions and are thought to be commensalisms in which seedlings benefit from the micro‐environment created by adult plants with no effect for the latter. This phenomenon is thought to be more frequent in harsh than in mild environments and appears to occur frequently in deserts and arid and semi‐arid biomes. Here, we investigate whether nurse‐protégé interactions are more common in arid environments by searching the published literature from the previous 92 years using the terms nurse plants, protégé plants, facilitation, nucleation and facultative mutualism. We then quantitatively compared these reports from arid zones to other environments. A total of 296 papers were found which referred to nurse‐protégé interactions. More than half (158) focused on arid and semi‐arid zones. This information was also used to explore hypotheses of potential causative forces that might have selected for such interactions in the arid zones such as seed trapping, nutrient, moisture, protection from browsing or trampling and support availability. Because of the large number of different nurse species (147, from 98 genera and 40 families) and protégé species (429, from 273 genera and 84 families), described across a diversity of environments, we suggest that there may be more than one causative factor selecting for nurse‐protégé interactions in arid and semi‐arid environments.     

Plant and small mammal richness correlate positively in a biodiversity hotspot

Hotspots of biodiversity are important areas in facilitating an understanding of species richness and its maintenance. Herbivores can increase plant richness by reducing dominant plant species thus providing space for subdominant species. As small mammals are abundant in the Succulent Karoo and therefore might affect plant richness by means of herbivory, we tested if this mechanism might exist in the Succulent Karoo in southern Africa, a biodiversity hotspot due to its extraordinary plant richness. At ten ecologically different study sites we measured plant and small mammal richness and diversity and determined 11 abiotic factors including soil composition, altitude and rainfall. We found positive correlations between plant richness and the number of small mammal species. A general linear model revealed that the number of small mammal species was more important than abiotic factors in explaining variation in plant richness. To test whether small mammals might directly influence plant richness, we studied the influence of the bush-Karoo rat Otomys unisulcatus, a central place forager, on the plant community. The immediate surroundings of occupied O. unisulcatus nests showed significantly higher plant richness than control areas. We conclude that small mammals can have a positive effect on plant richness in the Succulent Karoo. While experimental data are needed to support these correlative results, the results of our study indicate that areas of high small mammal richness should be included in conservation programs of the Succulent Karoo.     

Vulnerability of birds to contaminated water sources in the Karoo region of South Africa

The Karoo is a unique region in South Africa in terms of its ecological processes and endemic species. Large areas are needed to maintain viable populations of nomadic birds that follow erratic rainfall events and subsequent food and nesting resources, as well as ephemeral standing water. Whereas many species are adapted to arid conditions, our trait-based analysis found that an unusually large percentage (almost 45%) of 315 bird species in the semi-arid Karoo region rely on water to some degree. Indeed, some birds may have benefited from human activities to date, such as through the provision of water for livestock. However, this reliance on water makes birds vulnerable to changes in water quality stemming from various industrial developments. Given the large areas of the Karoo under consideration for concessions, the most noteworthy of these is hydraulic fracturing for shale gas, which results in a large quantity of waste water (‘produced water’) that contains a wide variety of chemicals, including petroleum byproducts. Given the negative impacts of secondary waste water on wildlife in other parts of the world where shale-gas exploration is being undertaken, careful attention must be given to preventing access to such produced water by an estimated 60 to 141 species of birds that make use of the water in the Karoo.