Ecosystems and Ecology Ecosystems and Ecology Author: Prof Koos Bothma Licensed under a Creative Commons Attribution license. CASE STUDY: THE SOUTH-WESTERN KALAHARI ECOSYSTEM The south-western Kalahari is a vast, arid ecosystem in southern Africa where ecological tolerances are stretched to the limit in a never-ending pursuit of survival and propagation by vegetation and animals. It will be used here as a case study to illustrate the concept of an ecosystem, although it is by no means an intricate one. This tumbleweed at sunset on a bare red dune near Loffiesdraai windmill in the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in August 1999 is the wind-blown, dried inflorescence messenger of the Bushman poison bulb Boophane disticha The Kalahari Sand System is the largest area of contiguous sand in the world and it underlies numerous ecosystems and bioregions. Its sands are discussed in some detail by Dougill and Thomas (2004) in the reference below. It stretches from the Orange River in South Africa north along eastern Namibia, western Botswana, western Zambia and eastern Angola to the tropical south-western parts of the Democratic Republic of the Congo, as far as 1°N. It underlies two great wetlands, the Okavango Delta of Botswana and the Liuwa Plain of western Zambia. Its south-western corner is most arid because the mean annual rainfall increases to the north-east. The south-western Kalahari is a semi-desert that forms a separate ecosystem within the Savanna Biome and it contains four distinct bioregions. These bioregions contain seven large landscape classes and 20 different landscapes based on morphology, soil and vegetation. The seven large landscape classes are: the high dunes; the low dunes; the plains on deep, reddish sand; 1|Page Ecosystems and Ecology the plains and terraces adjacent to the riverbeds; the grassy pans on compact whitish calcrete outcrops; the scattered patches of irregular dunes; and the riverbeds and pans. The term Kalahari was probably first used by the naturalist explorer A. Smith for this region in 1834 and refers to the Kgalagadi people who then inhabited it. The name Kgalagadi means place of thirst. Today the south-western Kalahari forms an integral part of the Kgalagadi Transfrontier Park of some 37 991 km² following the amalgamation of the Kalahari Gemsbok National Park of South Africa with the adjacent Gemsbok National Park of Botswana on 7 April 1999. The official launch took place on 12 May 2000. Inorganic components The major inorganic components of the south-western Kalahari ecosystem are its geology, soils, climate, and water resources. Geological formations The thick bed of sand that covers most of the surface of this ecosystem makes it difficult to study the underlying geology, most of which is known only from borehole cores. The rocks below the sands are part of the Karoo Sequence which consists of the Dwyka and Prince Albert Formations with some dolerite intrusions as vertical dykes. The Dwyka Formation was deposited during an ice age and consists of pebbles within a fine shale-like matrix. The Prince Albert Formation consists of shale that was deposited in an ancient shallow sea. The pre-Kalahari surface was dissected by rivers which drained north-eastward towards Botswana. The clayey gravel Wessels Formation and its overlaying clayey Budin Formation originated from Karoo Sequence rocks while the overlaying sandstone, grit and conglomerate of the Eden Formation yielded most of the sand. All these Formations were deposited under conditions that were wetter than present. The later calcrete of the Mokalanen Formation and sand of the Gordonia Formation indicate a change to a more arid environment while the clayey, diatomaceous limestone of the Lonely Formation was deposited in erstwhile lakes during a wetter phase in an otherwise arid period. The clay and sand of the Goeboe Goeboe Formation were deposited by water flow into the now normally dry pans and riverbeds. The Gordonia Formation represents the wind-blown sands that currently cover much of the land surface. There are no surface outcrops of the Wessel and Budin Formations. The Wessel Formation consists of deeply deposited gravels that were probably derived from glacial tillite deposits in the Dwyka Formation. The Budin Formation consists of red clay which came from the Dwyka and Prince Albert Formations and accumulated in a pre-Kalahari riverbed. The Eden Formation is visible on the surface along the current riverbeds, especially in the Auob River Valley. This Formation was probably deposited as a braided river system on a low relief surface and consists of visible but variable red, brown, yellow, cream and green sandstones, grits and conglomerate 2|Page Ecosystems and Ecology lenses. The sandstone sediments, that are less well consolidated than the rest, contain a network of hollow tubes that were either created by burrowing organisms or plant roots. The Mokalanen Formation consists of nodules which are hardpan calcrete outcrops that were deposited in an already semi-arid environment. The Lonely Formation occurs as clayey, diatomaceous limestone steps along the sides of some of the larger pans. These calcrete deposits contain the fossilized skeletal remains of diatoms and gastropods (molluscs) and have been intruded by the roots of the current vegetation. These fossils are always associated with pans and water courses and indicate that this Formation was formed in a shallow pan or a sluggishly flowing river with some aquatic vegetation. This probably happened during a wetter phase which occurred some 19 000 to 12 000 years ago in what was a long-term arid period when windblown sand was already present. The Goeboe Goeboe Formation occurs as horizontal sediments on the pans and riverbed floors. These sediments are relatively young and some are still forming at present. The main components are quartz, calcite, montmorillonite and feldspar. The surface sands of the Gordonia Formation are the most abundant geological feature in this ecosystem. They consists of sand of varying colour depending on the degree of iron dioxide which is present. These sands contain the iron-rich minerals magnetite, haematite and ilmemite among other heavy minerals such as garnet, rutile, tourmaline and zircon. Geomorphology There is a variety of dunes and dune patterns that have formed on the surface. Near the riverbeds the dunes are simple and linear but they become partially transverse and tend toward barchanoid ridges further away. In isolated spots, irregular, ellipsoidal, parabolic dunes have formed where the surface of older, stabilized dunes have become mobile again following the loss of vegetation and wind action. In other parts dunes are less developed while there are two major riverbeds. These rivers seldom flow and only do so following excessive rainfall in their catchments areas in Namibia. The Auob River flows more regularly than the Nossob River. Without its sandy surface cover and with the current arid climate the south-western Kalahari would be a young plateau where vertical erosion by the rivers that cross it has ceased. Some of the pans along the Nossob Riverbed were formed when the flow of water could no longer remove the sand load rapidly enough. The pans away from the rivers have become deflated by wind action and trampling by larger wildlife. 3|Page Ecosystems and Ecology The most arid south-western part of the south-western Kalahari ecosystem is covered by extensive parallel red dunes that are interspersed by well-vegetated interdune valleys as is shown south of the Auob River in this aerial view at sunrise in the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in August 1999 Climate The south-western Kalahari developed after the dissection of the topography during the wetter Miocene period which lasted from some 24 to 7 million years ago when the then existing river valleys were filled with sandy clay. Parts of these sediments were then re-deposited as poorly rounded and sorted conglomerates. When an arid period started some 7 million years ago the calcrete and silcrete deposits and outcrops were formed. Another wetter period then caused the onset of active erosion by rivers that now drained towards the south. The most recent wet phase, which lasted from some 19 000 to 12 000 years ago, allowed diatoms and molluscs to flourish in the pans. A subsequent arid period and the prevailing winds then formed most of the linear dunes and filled parts of the Nossob riverbed with deep sediments. Currently the ecosystem is experiencing a relatively wetter period which has stabilized the sand and vegetation. 4|Page Ecosystems and Ecology Sociable weaver birds Philatairus socius are typical of the south-western Kalahari ecosystem where they nest in dead trees such as this one near Dankbaar windmill in the Kgalagadi Transfrontier Park in June 1982 The current wide ambient temperature range and erratic rainfall create an annual hot, dry season, a hot, wet season and a cold dry season. The erratic annual rainfall usually varies from 185 to 230 mm, but some years are exceedingly dry and others exceedingly wet. The month of highest rainfall is January and the annual rainfall is cyclic with nine years of high rainfall followed by nine years of low rainfall. The mean annual rainfall increases north-eastward into Botswana with a low mean of 150 mm per year in the extreme south-western part of the south-western Kalahari ecosystem. The ambient temperature varies from -10.3° C in the winter to 45,4° C in the summer, but open sand surfaces will daily reach a peak temperature of 70° C or more in the summer. Light snow in midwinter has been recorded but is a rare phenomenon. In the winter ice may rarely form on the dune surfaces when they are wet. Soils The aeolian soils of the south-western Kalahari are broadly called Kalahari sands. However, there are five distinct groups of Kalahari sand, with the red and yellow sands being dominant. These soils support the terrestrial organisms and they anchor the plants. Through colloidal clay-humus complexes the soils also provide solutions from which the plants obtain water and mineral nutrients and provide a habitat for the micro-organisms that are essential in the decomposition of organic matter that recycle vital biochemicals. The red and yellow Kalahari sands form the parent material of the red Hutton and yellow Clovelly soil forms. These soils vary in thickness from a few centimetres to several metres or deeper. Soils near the riverbeds contain limestone materials which vary in colour from white to yellowish-white and grey. The uppermost layer of these soils is often hard and impervious and breaks into blocks 5|Page Ecosystems and Ecology of boulder limestone. Soils in the pans contain calcareous materials and soft, powdery lime that absorbs water well. A tumbleweed; the windblown, dried inflorescence of the Bushman poison bulb Boophane disticha; that has become lodged near a silver cluster-leaf tree Terminalia sericea near Loffiesdraai windmill in the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem in January 1997. These trees only grow on areas with deep sand The main Kalahari sand groups are red eutrophic, fine and medium sandy soils of the Hutton soil form; yellow-brown eutrophic and calcareous fine and medium sandy soils of the Clovelly soil form; deep calcareous reddish-brown and grey-brown loam and clay loam soils of the Oakleaf, Dundee and Valsrivier soil forms; brown calcareous sandy clay loam soils of the Swartland soil form; and shallow brown and yellow-brown calcareous sands and loams of the Mispah soil form. The Hutton soil form mainly gives rise to the typical red dunes of the south-western Kalahari. These sands are largely aeolion (wind-blown) in nature and the quartz grain surfaces are covered in iron oxide. The quartz grains are well-rounded and sorted as a result of the wind action. These homogeneous sands are freely permeable to water and absorb most of the rainfall rapidly to store moisture deeply, with a good grass cover developing in consequence. This moisture lasts well into the winter and the sands are penetrated on the surface by burrowing animals, such as gerbils, golden moles and mole-rats. The Clovelly soil form is similar to the Hutton soil form in its aeolian origin and the texture of its sands. However, the quartz grains have a yellow to yellow-brown colour due to different chemical and mineralogical properties. The soils are eutrophic with little horizon differentiation and therefore have a high exchange of calcium, magnesium, sodium and potassium. Some of these soils occur as undulations and dune-like formations on the banks of and in the immediate vicinity of the Auob and Nossob riverbeds. Others occur on the eastern, leeward side of pans and 6|Page Ecosystems and Ecology deflected basins. These sands are freely permeable to water but are underlain by an impermeable calcrete substratum. The surface sands form a habitat for some plants and for burrowing animals such as aardvarks, porcupines, springhares, mole-rats, mongooses, honeybadgers and a host of mice and rats. Brant’s whistling rat Parotomys brantsii is a typical, diurnal, rodent that burrows extensively in the south-western Kalahari ecosystem and feeds on the succulent stems of grasses and the leaves of bushes. This one was photographed in the Kgalagadi Transfrontier Park in August 1999 The Oakleaf, Dundee and Valsrivier soil forms are confined to the beds and floodplains of the Auob and Nossob Rivers and occur as narrow but continuous strips. The soils are alluvial in origin and often consist of loam or clay loam. They vary in colour from reddish-brown to brown and grey-brown and commonly contain lime concretions and nodules in the subsoil. These soils are alkaline with high concentrations of soluble salts. Their origin is mainly Karoo sediment rocks and mica with poor water penetration. The mica originated in the catchment areas in Namibia and is abundant in the Nossob River. These soils become impermeable when wetted and have a low productivity. Soils of the Swartland soil form are the most important component of the pans. They occur as thin, light brown topsoil with an extremely hard consistency and are highly calcareous and alkaline. In terms of plant producers the soils of this ecosystem are well supplied with the essential nutrients for plant growth and they are relatively free from the effect of leaching. However, the soils are generally low in nitrogen while the Hutton soil form is also low in phosphate. Cattle that graze the available require phosphate licks. The soils of the pans and riverbeds are more fertile than those of the dunes as they have higher phosphate levels. Consequently these pans and riverbeds form an important habitat for wildlife. 7|Page Ecosystems and Ecology The pans and riverbeds are normally dry and the rivers flow infrequently at intervals of several decades. The pans away from the riverbeds require self-maintenance because they are not subject to even occasional flooding from other regions and the combined interaction of their biotic and abiotic components is essential in these processes. Artificial waterholes may be highly mineralized and many contain high concentrations of toxic elements such as arsenic, mercury and nitrates (see Bothma 2005 below). The pans also form an important source of essential minerals for the larger herbivores. Antelopes such as this gemsbok Oryx gazella in the dry Nossob riverbed of the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in July 2001 eat the soil of pans and other brackish areas to supplement mineral deficiencies such as salt in their diet Organic components The main environmental factor that effects the production of the organic trophic levels in the southwestern Kalahari ecosystem is sufficient rainfall. It has been shown that even the denuded destruction zone around a waterhole will be repopulated by the best possible grazing following abundant rainfall. Drought and rainfall periodicity therefore have a significant impact on the herbaceous plants, but the deep-rooted woody plants are able to survive long periods of minimal rainfall. Vegetation The vegetation of the south-western Kalahari can be placed into 20 landscapes, which are the focal points for its large animals. These landscapes were described with the aid of Landsat ETM satellite images and 1:5000 orthophoto maps and earlier vegetation and habitat descriptions. There are seven large landscape groups in the south-western Kalahari ecosystem which contain the following landscapes: 8|Page Ecosystems and Ecology The parallel, grass-covered, high dunes near the banks of the Auob River mainly contain the grey camel thorn tree Acacia haematoxylon as a woody plant and are dominated by the tall dune bushman grass Stipagrostis amabilis. Prostrate creepers such as the moisture-rich gemsbok cucumber Acanthosicyos naudianus and tsamma melon Citrullus lanatus occur seasonally on the naked dune crests and are an important water resource for wildlife. The tsamma melon Citrullus lanatus of the south-western Kalahari ecosystem is a major source of moisture for wildlife and the San people. Tsamma melons, such as these near Loffiesdraai in the Kgalagadi Transfrontier Park in January 1997, consist of up to 90 per cent water The irregular high duneveld occurs south of the Auob River and between the Auob and Nossob Rivers in the south. It contains scattered grey camel thorn trees, the black thorn Acacia mellifera and the shepherd’s tree Boscia albitrunca as a shrub, the latter acting as ideal cover for the larger carnivores. The grasses vary but the three-thorn bush Rhigozum trichotomum dominates the calcareous soils in the valleys between the dunes and its leaves and flowers are an important source of browse. Mainly close to but north of the Auob River and on both sides adjacent to the Nossob River the parallel high dunes support an open tree savanna mainly consisting of the camel thorn tree Acacia erioloba and the black thorn tree, with a variety of other woody shrubs and grasses. This landscape forms an important grazing area for wildlife that move there from the riverbeds at times. The low dunes mainly occur in the interior areas away from the riverbeds. A combination of the camel thorn tree and the mostly unpalatable Kalahari sour grass Schmidtia kalahariensis forms an open tree savanna in the northern part of the ecosystem. This landscape contains numerous herbivores such as springbok, blue wildebeest and gemsbok, and carnivores such as lions, cheetahs, spotted hyaenas and black-backed jackals. 9|Page Ecosystems and Ecology The riverbed of the fossil Nossob River, as seen here at Lijersdraai in the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in January 1984, has an abundance of sweet grazing during the rainy season The irregular low dunes are a large landscape of grass-covered irregularly shaped dunes in the more arid southwestern part of this ecosystem north of the Auob River. It contains numerous large pans and the grey camel thorn tree is the most conspicuous component of the woody vegetation. There is a large variety of grasses and forbs. The vegetation of the dunes differs from that of the interdune valleys with the black thorn tree, the shepherd’s tree and the grey camel thorn tree being the most conspicuous woody plants. The interdune valleys mainly contain the grey camel thorn tree, the blackthorn and the three-thorn bush. The second large landscape of this ecosystem consists of the undulating low dunes that occur throughout the area between the Auob and the Nossob Riverbeds. The prominent woody plants are the camel thorn tree, the grey camel thorn tree and the shepherd’s tree. The landscape contains several palatable species of grass and various types of dwarf shrub. There are two types of plains landscape on reddish sand. Grey camel thorn trees and dune bushman grass occur in large patches throughout the interior regions away from the riverbeds but the dunes are seldom more than 6 m high. Other palatable grass species attract large herbivores to this landscape. The flat grassy plains with scattered shrubs that also occur on reddish sand in the central interior region are slightly undulated and contain scattered but low grey camel thorn trees. The calcrete outcrops, terraces and floodplains along the Auob and Nossob Rivers form an important wildlife habitat. They contain scattered individual camel thorn, grey camel thorn and shepherd’s trees and some dwarf woody shrubs but the grass layer is poorly developed. There are four types of landscape on whitish sand on the plains and terraces adjacent to the riverbeds and grassy pans. The scattered large camel thorn tree, three-thorn bushes and small 10 | P a g e Ecosystems and Ecology bushman grass Stipagrostis obtusa clumps occur on slightly compact calcareous and whitish sand in the dune valleys, near pans and adjacent to the Auob and Nossob Rivers and provide shade and palatable forage to large herbivores in the wet season. After rains these flats are covered in yellow beds of the devil thorn Tribulus zeyheri. The black thorn, three-thorn and small bushman grass landscape of the riverbed terraces occurs on the calcareous terraces near pans and the rivers and also provides good grazing to short-grass grazers such as blue wildebeest in the wet season. A small landscape unit of dwarf ganna shrubs Solsola spp, mixed with the kapok bush Eriocephalus spp., occurs on the edges of the pans and calcareous outcrops adjacent to the Nossob River, while large patches of uniform small bushman grass form a further landscape unit in local patches on whitish, compact sandy soils on terraces and floodplains next to the Nossob River. The sap of the bulb of the Bushman poison bulb plant Boophane disticha as shown here in the interior sandveld dunes of the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in June 1997, is used as a toxin in the poison arrows of the San people when hunting The riverbeds and pans contain six types of landscape. The pans both have bare floor surfaces and are commonly known as salt pans, or they have grassy surfaces. Both types occur on whitish, strongly alkaline calcareous and silted soils. The vegetation on and around a pan differs markedly from that of the surrounding area and often shows concentric zoning. Around some of the larger pans in the northern part of this ecosystem, stands of the worm-bark false-thorn tree Albizzia anthelmintica occur. The riverbeds contain four types of landscape with a dense layer of white buffalo grass Panicum coloratum and the scattered large camel thorn trees in the northern part of the Nossob riverbed. This grass offers good grazing to the larger antelopes. An isolated stand of the blue bush Lebeckia linearifolia occurs in the lower Auob and Nossob Riverbeds and on some high parallel dunes in the vicinity. There is no grass cover in this landscape unit. From its confluence with the Auob River northward for some 160 km a landscape that is characterized by shrubs and scattered camel thorn trees with a limited grass cover occurs. The Auob Riverbed represents a narrow zone of large camel thorn and grey camel thorn trees on moderately 11 | P a g e Ecosystems and Ecology compacted calcareous sandy soils while the dominant types of grass are the unpalatable sour Kalahari grass and the palatable small bushman grass. The final landscape is the striking and highly irregular dune landscape with its striking ellipsoidal parabolic dunes which are only found in a few isolated places within the interior duneveld. The only woody plants in this landscape unit are a few black thorn and shepherd’s trees and shrubs. Floristically the south-western Kalahari ecosystem is impoverished with a reduced landscape differentiation. The camel thorn tree, the grey camel thorn tree and the shepherd’s tree are characteristic of several types of landscape and are all listed as protected trees in the National Forests Act of South Africa (Act 84 of 1998). The camel thorn tree has a significant ecological role for numerous types of animal for feeding, nesting or resting. Grazing impacts are localized by a natural rotational grazing system that involves localized and long-distance movements by the major herbivores, often following rainfall. The interior dunes and plains are only utilized by wildlife during the wet season. Geology, soil and climate have therefore dictated the nature and development of the vegetation of the south-western Kalahari. In turn the vegetation dictates the number, distribution and type of larger herbivores that can be sustained there. Consequently it also dictates the number and distribution of those larger carnivores that prey on these herbivores. The cheetah Acinonyx jubatus often utilises the open Nossob riverbed of the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem, as seen here in January 1975, as a hunting ground because these open plains do not impede its high-speed chases Animals The south-western Kalahari ecosystem is the realm of a surprisingly rich invertebrate fauna about which little is known. The invertebrate faunal complement includes 37 genera and 51 species of sand-adapted tenebrionid beetle which live in the dunes and form 60 per cent of all the beetles of the south-western Kalahari. 12 | P a g e Ecosystems and Ecology Of the vertebrates, the reptiles all have arid zone affinities and this ecosystem coincides with a number of distributional range limits. Endemic taxa include the gecko Colopus wahlbergi furcifer and the Kalahari serrated tortoise Psammobates oculifer. The poorly known blind snake Typhlops boylei also occurs. There is also a range of amphibians, with the bushveld rain frog Breviceps adspersus adspersus, the giant bull frog Pyxicephalus adspersus adspersus and the Tremelo sand frog Tomopterna cryptotis reaching their southernmost distribution limit in this ecosystem. Snakes such as this puff-adder Bitis arietans at Gharagab windmill in the Kgalagadi Transfrontier Park in January 1978 are abundant in the south-western Kalahari ecosystem and are especially deadly to cats such as the cheetah, leopard and lion The birds of the south-western Kalahari are sedentary, insectivorous or carnivorous, and nongregarious; or they are nomadic granivores that are gregarious even when they are breeding. All the ground-dwelling sedentary birds are cryptically coloured to avoid predation. Breeding is initiated in most species by the onset of the wet season or by associated ecological effects. Nest orientation is related to the season of breeding and capitalizes on maximal shade in the hot season. Evaporative cooling is employed by all the birds of this ecosystem where surface water is a rare commodity. The use of large communal nests by the sociable weaver Philetairus socius is also a thermoregulatory strategy. The ostrich Struthio camelus is a large terrestrial and herbivorous bird. 13 | P a g e Ecosystems and Ecology The huge sociable weaver Philatairus socius nest in a dead camel-thorn Acacia erioloba tree near Ardlamont windmill in the Kgalagadi Transfrontier Park in the south-western Kalahari ecosystem in 1991 could contain op to 300 nest chambers which are occupied by a pair of birds each This male ostrich Struthio camelus in the Nossob riverbed in the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem in July 2001 represents the largest, living terrestrial bird in the world Much of the plant production in the south-western Kalahari is utilized by the rodent complement whose populations fluctuate greatly with the amount of rainfall received and therefore the amount of plant food being produced. In turn this affects the occurrence and abundance of raptors such 14 | P a g e Ecosystems and Ecology as the tawny eagle Aquila rapax, martial eagle Polemaetus billicosus, bataleur Terathopius ecaudatus and southern pale chanting goshawk Melierax canorus, and the smaller mammal carnivores such as the Cape silver fox Vulpes chama, black-backed jackal Canis mesomelas and caracal Caracal caracal which mainly feed on the rodents. There are 17 species of rodent that are known to occur in the south-western Kalahari ecosystem, with their sizes varying from as little as 5 g to 3 kg, with the majority weighing less than 100 g. Most of the rodents are diurnal but some activity will continue on moonlit nights. Only the common mole-rat Cryptomys hottentotus is totally fossorial. Most of the rodents are granivorous but some are also partially insectivorous in their diet. The rodent mole and springhare Pedetes capensis eat a variety of plant material. Major fluctuations in the rodent population occur mainly in response to rainfall fluctuations. The hairyfooted gerbil Gerbillurus paeba and the four-striped grass mouse Rhabdomys pumilio are, however, consistently numerous and support the maintenance of a minimum raptor and mammal carnivore population. Most of the reproductive activity happens in the hot, wet season. The lack of large diversity in the rodent population is probably caused by the absence of particular habitat requirements although the species diversity in this ecosystem is larger than expected. The tree rat Thallomys paedulcus, photographed in January 1985 at Loffiesdraai windmill in the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem, utilises the camel-thorn tree Acacia erioloba as its habitat where it feeds nocturnally on the fine leaves. These trees form a habitat and source of food for a variety of animals The south-western Kalahari ecosystem contains a relatively small population of smaller carnivores. The yellow mongoose Cynictis penicillata is common but prefers a dune habitat while the suricate Suricata suricatta prefers a riverbed one. The other species show no clear habitat preference. The small black-footed cat Felis nigripes is rare as this ecosystem is at the western end of its distributional range. The only other small cats that occur in the south-western Kalahari are the wildcat Felis sylvestris and the caracal. The slender mongoose Galerella sanguinea is encountered occasionally along the riverbeds while the banded mongoose Mungos mungo is known only from rare sightings. Invertebrates, reptiles, rodents, hares, birds and smaller antelope form the main diet for the wildcat and the caracal. Foraging honey badgers Mellivora capensis are 15 | P a g e Ecosystems and Ecology frequently accompanied by chanting goshawks and occasionally also by black-backed jackals. Many rodents that are dug out of their burrows by the badgers and manage to escape capture by the badgers are then captured by the goshawks and jackals in a form of symbiotic hunting. The aardwolf Proteles cristatus is a specialized carnivore that almost entirely feeds on long-nosed harvester termites of the genus Trinervitermes. This suricate Suricata suricatta attacking a scorpion at Twee Rivieren in the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem in January 1989 is a small, diurnal, burrowing and sociable carnivore that prefers the riverbeds as habitat and mainly eats invertebrates The canids are represented by three medium-sized species which includes bat-eared fox Otocyon megalotis, Cape silver fox and black-backed jackals. Their diet mainly consists of smaller animals, with the bat-eared fox being mainly insectivorous in diet. In the south-western Kalahari ecosystem Cape silver fox and black-backed jackal are not known to attack the lambs of smaller antelope and their largest prey seems to be the springhare. In the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem, the bat-eared fox Otocyon megalotis as shown here in the Auob riverbed in July 1981, is a nocturnal insectivorous carnivore 16 | P a g e Ecosystems and Ecology There are three types of hyaena in the south-western Kalahari ecosystem, with the aardwolf specializing on long-nosed harvester termites, the brown hyaena Hyaena brunnea being mainly a scavenger and the spotted hyaena Crocuta crocuta a scavenger and an adept hunter when hunting as a pack. Brown hyaenas will also eat wild fruits, including the moisture-rich tsamma melon, when they are available. The spotted hyaena is dependent on a regular surface water supply. The larger cats include lions Panthera leo, leopards Panthera pardus and cheetah Acinonyx jubatus and they all hunt mammal prey opportunistically. Consequently, their survival depends largely on the abundance of ungulates such as the gemsbok, blue wildebeest, eland, red hartebeest, springbok, grey duiker and steenbok which in turn is dependent on the abundance of rainfall and the resultant vegetation quantity and quality. These herbivores move locally and regionally in response to rainfall and will suddenly congregate in large numbers in the riverbeds after suitable abundant rain. The provision of artificial waterholes has, however, caused some of the herbivore population to become sedentary. In times of drought the larger carnivores will readily make use of other prey such as the smaller carnivores, porcupines Hystrix africaeaustralis, the smaller and more sedentary antelope and even springhares. Cheetah seldom take prey larger than springbok Antidorcas marsupialis and are mainly found along the riverbeds which these antelope favour and which provides an ideal hunting ground for a high-speed chase. Leopard and lion utilize the larger herbivores such as the eland, gemsbok and blue wildebeest too but they are opportunistic hunters and will prey on whatever suitable prey becomes available. The lion is the apex predator in this ecosystem. The fossil riverbeds such as that of the Nossob River at Kwang in the Kgalagadi Transfrontier Park of the south-western Kalahari ecosystem contain many ancient loops and ox-bows where lions Panthera leo may find shelter in the shade on a hot day such as in January 1984 17 | P a g e Ecosystems and Ecology Conclusions The south-western Kalahari ecosystem is relatively waterless with erratic and low rainfall. It is a largely featureless and marginal habitat and not as complex as some other ecosystems. Nevertheless it consists of a matrix of food chains of a tenuous nature. Any disturbance in one element is likely to create ripple effects throughout the ecosystem. The best management approach seems to be to have as little as possible human intervention. The provision of artificial water may be the only intervention that could increase the ecological capacity of this ecosystem for wildlife. Despite appearing to be ancient, the current ecosystem seems to have developed relatively recently following an extensive wet period that ended some 12 000 years ago, although it is currently experiencing a wetter phase within a dry period. If and when environmental conditions change again, the current delicate equilibrium is likely to shift as thresholds of tolerance are passed. 18 | P a g e