Trophic status of small mammals in the semi-arid Karoo, South Africa

The feeding habits of a community of small mammals from the semi-arid Karoo, South Africa were analysed by microscopic examination of the stomach contents of animals caught during a 13-month snap-trapping study. The community comprised eight rodent species ( Gerbillurus paeba, Mus minutoides, Rhabdomys pumilio, Otomys unisulcatus, Saccostomus campestris, Desmodillus auricularis, Malacothrix typica, Mastomys natalensis ) and a single macroscelid elephant shrew ( Macroscelides proboscideus. ) The rodents were all predominantly herbivorous, while the elephant shrew ate mainly insects. No granivores or omnivores were represented in this community. The results presented here (including the first analysis of the diet of M. typica ) indicate a greater emphasis on herbivory than previously described for these species. Limited dietary overlap was recorded between these species, although some instances of potential dietary competition that deserve further study were identified. The paucity of granivorous small mammals in the semi-arid Karoo contrasts the patterns of trophic specialization for North American and Israeli semi-arid communities, but is similar to that of South American, Australian and other southern African semi-arid communities.     

Disturbances in semi-arid shrubland and arid grassland in the Karoo, South Africa: mammal diggings as germination sites

Mammals (mainly aardvark Orycteropus ufer , porcupine Hystrix austro-africanae , bat-eared fox Otocyon megalotis and Cape fox Vulpes chama ) which excavate holes in hard-capped soils in Karoo shrubland and inter-dune pans create microsites where seeds, detritus and water accumulate, and seeds germinate. Holes with seedlings were significantly deeper (6.7 ± 0.4 cm) than holes without seedlings (4.5 ± 0.3 cm). In open, inter-shrub sites, significantly more seedlings germinated in diggings than on the flat soil surface, but there was no difference in the proportion of seedlings germinating under shrubs and in diggings. In inter-dune pans, 96% of all seedlings found were growing in diggings.

Résumé

Les mammifères (surtout l'oryctérope Orycreropus afer , le porcépic Hystrix austro-ajricanae , l'otocyon Otocyion megalotis , et le renard du Cap Vulpes chama ) qui creusent des trous dans les régions buissonneuses du Karoo et dans les creux entre les dunes créent des microsites où les semences, les déchets et l'eau se rassemblent, et les semences germent. Les trous où elles germent sont significativement plus profonds (6,7 ± 0,4 cm) que ceux où elles ne germent pas (4,5± 0,3 cm). Aux endroits buissonneux ouverts, il pousse plus de jeunes plantes dans des creux qu'à la surface du sol, mais il n'y a pas de différence entre les proportions de celles qui poussent sous les buissons et dans les creux. Dans les creux entre les dunes, 96% des jeunes pousses croissent dans les excavations.     

Small mammal seed consumption in the Karoo,South Africa: further evidence for divergence in desert biotic processes

Summary Annual seed consumption by a small mammal community in the semi-arid Karoo, South Africa, was calculated using population, biomass and dietary estimates and published estimates of field metabolic rate. The community comprised a macroscelid elephant shrew Macroscelides proboscideus, and two rodents Gerbillurus paeba and Desmodillus auricularis. Only G. paeba consumed seed during the study, and then in low amounts (annual mean = 0.4% of diet, insects = 53%, herbage = 47%), with M. proboscideus and D. auriocularis consuming mainly insects (88%) and herbage (90%), respectively. Energy requirements for the G. paeba population averaged 568 kJ ha^–1 day^–1. Total seed consumption was estimated to be 140 g ha^–1, or 155 000 seeds ha^–1 which represents 0.5% of the annual seed production for this site. These levels of granivory by small mammals are significantly lower than those recorded for North American deserts, and raises the question as to why granivory varies between these systems. These findings also refute the hypothesis that as a result of the harshness and similarity of physical conditions in deserts, the properties of desert communities are convergent.     

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.     

Changes in woody community structure and composition under constrasting landuse systems in a semi-arid savanna, South Africa

We aimed to explore the farm scale effects of three landuse types, communal grazing, wildlife management and commercial cattle farming, on the woody vegetation of a semiarid savanna. Location The study farms were located within a single bioclimatic zone in semiarid savanna, South Africa. Methods The species composition and structure of woody vegetation on three farms of each of three landuse types were sampled. Results We found that communal grazing land sites were classified outside the topland-bottomland vegetation dichotomy characteristic of this region. Comparisons of size class distributions showed the communal grazing lands had fewer small and large individuals; suggesting both lower levels of regeneration and regenerative capacity in the communal grazing lands. The species richness and biomass of woody plants was lower on communal grazing lands than on private game reserves and commercial cattle farms. The longevity of tree species explains the observed lag between changes in abundance and species loss; we consequently predict that there will be future losses of species in the communal grazing lands. By classifying species into a range of use-categories we showed that utilization and species loss was not limited to certain plant use categories. Higher levels of wood harvesting measured in the communal grazing lands are likely to be responsible for the observed differences. Main conclusions It is concluded that communal grazing management at this study site has substantially changed the composition and structure of woody plant communities, and that these changes have reduced the current availability of natural resources and will reduce resource production in the future.     

Live fast,die young: estimating size-age relations and mortality pattern of shrubs species in the semi-arid Karoo,South Africa

We present a technique for estimating size-age relations and size-dependent mortality patterns of long-lived plants. The technique requires two sets of size data of individual (non-marked) plants that should be collected with a time-lag of several years in the same area of a study site. The basic idea of our technique is to assume general (three parameter) families of size-dependent functions which describe growth and mortality that occurred between the two data gathering events. We apply these growth and mortality functions to the size data of the early data set and construct predicted size-class distributions to compare it, in a systematic way, to the size-class distribution of the later data set. In a next step we calculate the size-age relations from the resulting growth functions, which yield the smallest difference between observed and predicted size-class distribution. Applying this technique to size data of five dominant shrub species at the Tierberg study site in the semiarid Karoo, South Africa produced new insight into the biology of these species which otherwise cannot be obtained without frequent measurements of marked plants. We could relate characteristics of growth behavior and mortality, for certain subgroups of the five species, to the life-history attributes evergreen vs. deciduous, succulent vs. woody, and early reproductive vs. late reproductive. The results of our pilot-study suggest a broad applicability of our technique to other shrublands of the world. This requires at least one older record of (individual) shrub-size data and performance of resampling.     

Environmental History and the Desertification of the Karoo,South Africa

Abstract

This paper reviews the desertification debate in South Africa and emphasizes the methods that have been used to understand the environmental history of the semi–arid rangelands in the eastern Karoo during the last 500 years. These mixed grass/dwarf shrub rangelands, with mean annual rainfall totals typically between 300–400 mm, are described and the main driving variables discussed. A brief historical account of the European settlement of the region is also presented. The desertification debate has focussed on three main issues. Firstly, because arid and semi–arid lands appear to be heavily influenced by climate, the changing climatic regime during the Holocene and particularly during the last two hundred years has been the subject of intensive investigation. While all studies conclude that there is no evidence in the historical record to support the popular perception that rainfall totals have decreased this century, the question of changing rainfall seasonality has not been adequately explored. Although mean annual temperatures over the last 50 years have not changed there have been significant increases in mean monthly maximum temperatures and significant decreases in mean monthly minimum temperatures for some of the stations investigated. The second issue of great interest in the South African desertification literature is that of the nature of pre–colonial environments. A wide variety of archaeological, historical and ecological techniques have been used, including analyses of fossil mammal bones in owl pellets, fossil pollen in hyrax middens, notes from traveller's accounts of the region and stable carbon isotopes in soil as well as fossil mammal bones. While all authors agree that the eastern Karoo was more grassy at some stage in the past there is disagreement as to both the timing and cause of the changes to a more shrubby vegetation. The final issue of great concern to the desertification debate in South Africa concerns the rate of change during the last 100 years. Satellite imagery, matched ground and aerial photography, survey data and an analysis of historical stock records cannot agree as to whether the Karoo is degrading or not. Certainly, the classic view of an annually expanding desert margin has been replaced in recent years by a more realistic understanding of the seasonal dynamics of the vegetation. The recent trend to detailed modelling of the demographic process in key species holds much promise for our understanding of the degradation process. The vibrant community of researchers, employing a range of archaeological, historical and ecological techniques, will make important contributions to South Africa's National Action Plan to Combat Desertification.     

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.     

Conservation value of road verges in the semi-arid Karoo, South Africa: ants (Hymenoptera: Formicidae) as bio-indicators

Ground-foraging ants (Hymenoptera: Formicidae) were used to assess the conservation value of road verges in a semi-arid region around Prince Albert, Western Cape Province, South Africa. Ant communities were sampled using pitfall traps on 50 sites along two roads. Four transects were sampled at each site, two in the road verge and two in adjacent rangeland. A total of 43,298 individual ants of 34 species were caught, with 31 and 32 species in the road verge and adjacent rangeland respectively, but on average road verges were the more species-rich. Road verges also contained relatively more rare species (i.e. those species that occurred in <10% of the traps. Ant species richness appeared to be influenced by food availability. Although ants appeared to be negatively impacted by higher grazing intensities in the rangeland, there was no difference in species richness between fenced and unfenced road verges. Species in the road verge may benefit directly from road kills, and indirectly from lower grazing pressure on plants, increased surface run-off and differences in soil surface temperatures. No alien ant species were observed or captured at any of the sites.     

Glacial cycling, large mammal community composition, and trophic adaptations in the Western Cape, South Africa

Some of the earliest evidence for modern human behavior has been recovered from the Western Cape Province, South Africa. Archaeological and paleontological sites in the Western Cape are typically described as “glacial” or “interglacial” in aspect based on the numbers of grazers found in the faunal assemblage, as glacial periods are often thought to have been characterized by spreading C₄ grasslands that replaced endemic C₃ shrubland vegetation found in the Western Cape today. Here, we test the hypothesis that glacial and interglacial time periods were associated with a predictable change in large mammal trophic adaptations by analyzing the proportions of grazing larger mammals from 118 levels of 15 Western Cape fossil assemblages sampling marine isotope stage (MIS) 6 to the present time to determine whether there is a change in composition in these communities that might reflect a shift in ecology and habitat. Our results indicate that trophic proportions did not significantly change over time in the Western Cape as a whole, and thus the hypothesis for habitat changes affecting the subsistence ecology of modern humans during the development of modern behavior is not supported. However, our results show that the southwestern subregion of the Western Cape was characterized by the presence of more grazing species through time than the western subregion. Thus, if ecological and population isolation during glacial periods were integral to catalyzing the development of modern behaviors in the Western Cape region of South Africa, then a complex model including the development of possible mosaic habitats is needed.     

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.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

Predictors of mammal species richness in KwaZulu-Natal,South Africa

The management of multi-functional landscapes warrants better knowledge of environment-richness associations at varying disturbance levels and habitat gradients. Intensive land-use patterns for agricultural purposes lead to fragmentation of natural habitat resulting in biodiversity loss that can be measured using landscape metrics to assess mammalian richness. Since carnivores and herbivores are likely to show different responses to disturbance, we calculated carnivore, non-carnivore, and total mammal species richness from camera surveys using a first order Jackknife Estimator. Richness was compared along a habitat gradient comprising coastal forest, Acacia thicket, and highland in KwaZulu-Natal, South Africa. We used standardized OLS regression models to identify climatic and disturbance variables, and landscape metrics as predictors of species richness. The estimated total and non-carnivore species richness were highest in coastal forest, while carnivore species richness was highest in highland followed by coastal forest and Acacia thicket. Average monthly maximum temperature was a significant predictor of all richness groups, and precipitation of the wettest month and isothermality determined total and non-carnivore species richness, respectively. These climatic variables possibly limit species distribution because of physiological tolerance of the species. Total mammal richness was determined by mean shape (+) and habitat division (−) while diversity (+) and patch richness (−) explained carnivore species richness. Mean shape index (+) influenced non-carnivore richness. However, habitat division and patch richness negatively influenced total mammal richness. Though habitat patch size and contiguity had a weak positive prediction, these metrics demonstrated the importance of habitat connectivity for maintaining mammal richness. The identification of these climatic and landscape patterns is important to facilitate future landscape management for mammal conservation in forest-mosaics.     

Physiological characteristics of geophytes in semi-arid Namaqualand,South Africa

Namaqualand, a semi-arid winter rainfall region of South Africa, supports an exceptional diversity of geophytic species. The survey focused on gas exchange reactions and chlorophyll a fluorescence in geophytes with different leaf orientation in relation to environmental variability. Although the above ground life cycle of geophytes can be extremely short, unlike desert annuals, they are not characterized by a high photosynthetic CO₂ uptake. Maximum CO₂ uptake ranged from 4 to 20 mol CO₂ m^-2 s^-1. Temperature optima of photosynthetic CO₂ uptake were comparably low and ranged from 12 to 22°C for eleven species tested, with only one species above 19°C. The decrease of CO₂ uptake with rising temperatures was associated with a substantial increase of photorespiration. Net photosynthesis was saturated between 500 and 900 mol photons m^-2 s^-1 while electron transport through PSII was saturated at higher photon flux densities. At light intensities beyond saturation, a high variability of PSII efficiency occurred. It was highest for horizontal leaves and lowest for upright leaves. However, the maximum quantum yield of PSII (F_v/F_m)remained constant during the course of a day, regardless of leaf orientation. This indicates the absence of photoinhibitory effects and a well protected photosynthetic apparatus. Leaf orientation determined interception of solar radiation and thus leaf temperature which was highest for horizontal leaves. In conclusion, Namaqualand geophytes show photosynthetic characteristics that are well adapted to the mild and generally moist conditions during the growing season.     

Influence of vegetation structure on the small mammal community in a shortgrass prairie ecosystem

The structure of vegetation, and how this structure varies across a landscape, is crucial to understanding the distribution of wildlife species. Between 2002 and 2004, we sampled small mammal communities and measured vegetation structure at 185 locations across a range of disturbance regimes in a shortgrass prairie ecosystem in southeastern Colorado, USA. At each sampling location, the local disturbance regime was some combination of varying intensity of livestock grazing, military training activity, and fire. Vegetation structural characteristics measured included percent bare ground, basal cover, litter, shrub density, and mean grass and shrub height. Rodent communities were described by richness, diversity, total and per capita biomass, and species abundances. Northern grasshopper mice (Onychomys leucogaster), Ord's kangaroo rats (Dipodomys ordii), silky pocket mice (Perognathus flavus), western harvest mice (Reithrodontomys megalotis), white-footed mice (Peromyscus leucopus), southern plains wood rats (Neotoma micropus), thirteen-lined ground squirrels (Spermophilus tridecemlineatus), deer mice (Peromyscus maniculatus), and spotted ground squirrels (Spermophilus spilosoma) accounted for >99 % of all captures. Canonical correlation analysis was used to assess the relationship between small mammals and vegetation structure. The first two canonical variates explained over 50 % of the variation in vegetation structure and were related to the ratio of bare ground to basal coverage and litter accumulation. Rodent community indices were most strongly related to litter accumulation and shrub density, though the models had low explanatory power. Our results agreed with published findings regarding microhabitat associations and indicated small mammal communities benefited from a system of interacting disturbances and the resulting landscape mosaic.     

The small mammal community of Mukogodo Forest, Kenya

Species richness and diversity of rodents and insectivores were investigated at relict forest patches of Mukogodo, Laikipia, Kenya using Sherman's live traps and pitfall traps. Two hundred and nineteen individuals were captured in 3021 trap‐nights. There were eleven species in two taxonomic groups, Rodentia and Insectivora. Two other rodent species were sighted but not captured. Thirteen bats belonging to four species (Epomophorous wahlbergi, Pipistrellus kuhlii, Scotophilus dingani and Nycteris thebaica) were opportunistically trapped using mist nets. Two of the four species accumulation curves for forest patches did not reach an asymptote. Species richness and diversity were highest at Kurikuri compared with other patches because of habitat variability. The results support the prediction that forest disturbance and degradation lead to an increase in generalist species as compared with specialists and highlight the importance of relict afromontane forests in the conservation of small mammals in Kenya.     

An updated plant checklist for Tankwa Karoo National Park,South Africa

An updated checklist is provided for the flora of Tankwa Karoo National Park, which occupies an area of 143,600 ha. A total of 730 species and 780 plant taxa (species, subspecies and varieties), representing 267 genera in 73 families, are recorded for the park. This comprises 30 bryophytes, 7 pteridophytes, 189 monocotyledons and 554 dicotyledons. Sixteen species are endemic or near-endemic to the Hantam-Tanqua-Roggeveld subregion. Twenty-eight species are of conservation concern and six species have been declared as alien invasives.