CHAPTER The Dynamics of Ecosystem Functioning oO U T C O M E § In the previous chapter it was shown that the environment consists of many elements that are functionally connected to each other in a dynamic way. After completion of this section, you should be able to: 3.1 Realize that these elements act in a very orderly way according to specific laws and guiding principles. 3.2 Understand food chains and energy flow through the ecosystem: 3.3. Discuss the continuous movement of elements through the system to ensure that ecosystem health and stability is retained. 3.4 Describe how population numbers are being kept in check by Nature. 3.5 Understand how dynamic balance is being retained in an ecosystem. 29P R E A M B L E We have for a long time been breaking the little laws, and the big laws are beginning to catch up with us. (AE Coventry) If you violate Nature's laws, you are your own prosecuting attorney, judge, jury and hangman. (Luther Burbank) 3.1 Natural laws “Pic oni for reason fo the foaming Wwironmental crisis” Is not simplistic. What is clear, Is that Nature in itself does not have a crisis. ‘The crises lie in human hands: people are experi- encing a major problem, namely, to adapt suc- cessfully to their habitat. Plants and animals fit into the system that function according to specific laws. Humankind has elevated itself above these laws and is paying the price. We will examine briefly some of the most relevant environmental laws before explaining the functioning of the environment as a system, As seen in the previous chapter, energy laws are fundamental to the functioning of the environmental system. 3.1.1 Thermodynamic laws TCOME Bal k You have to realize that the environmental factors act in a very orderly way according to specific laws and principles. ts ‘The environmental crisis is not fundamentally a de facto shortage of resources, but rather the inability of humans to muster sufficient energy to develop resources that are (relatively) abundantly available. Unlimited energy would (theoretically), for example, make it possible to change all the dry areas of the world into irrigated lands using desalinated water. The limitation of production therefore lies in the limitation of the thermo- dynamic laws. ‘The FIRST LAW states that there is a limit to available energy and that humans are unable to create energy independently (see par. 2.2.1.1). ‘The SECOND LAW states that the use of energy inevitably leads to entropy. This means that energy consumption results in a decline in its utility. Energy is therefore not re-usable. 30 6 Ecopian Environmental Management3.1.2 Preservation of matter ‘The law of preservation of matter restricts human's development, since matter, like energy, cannot be created. It can only be transformed. Implements can thus be manufactured from raw materials (elements) if there is sufficient energy. At present itis generally cheaper to mine raw materials and construct new products rather than 3.1.3 Limiting factors If a factor, e.g. the absence of a nutrient, pre~ vents or inhibits the establishment or growth of 2 species in an environment, itis known as @ limiting factor. In many cases the limiting factor is not @ macro-nutrient like nitrogen (N) or phosphorus (P) which is usually needed in large quantities, but rather essential micro-nutrients like boron (B) or molybdenite (Mo) which usually occur in very small quantities in the soil. In a similar way the growth of plants may be stunted Or inhibited as a result of an iron deficiency, because the pH is so low that it inhibits the uptake of iron (Fe). In order for a species to survive in a certain place the environmental conditions must be within the tolerance levels of that particular species. A species cannot survive below the lower, or above the highest tolerance limit (Fig. 3.1). Ifa factor, for example, moisture (rainfall) is plotted as a variable against the presence of a species, then the graph will show a normal dis- tribution curve. | itn et N \ oar WSS SN (Source: Rye, 1978) Figure 3.1: The normal tolerance curve of species distribution in terms of limiting factors re-cycle existing ones. Since production process- es are energy intensive and matter Is indestruc- tible, extensive waste generation and pollution ‘occur which in turn adversely affect the quality of the environment (see also par. 6.3.2.4 for a discussion of the application of these Laws to the environmental crisis). In other words, the species will occur most often in those areas where the moisture conditions are optimal. The occurrence will decrease in those areas where there is either too little or too much moisture, Those areas where the species can no longer survive are known as the bottom and top limits of tolerance. Fig. 3.2 indicates the distribution of the most important terrestrial biomes in terms of their temperature and moisture tolerance. The prin- ciple of limiting factors shows that where a limiting factor exists, for example the absence of a plant nutrient, which inhibits growth, and this nutrient is supplemented, the growth will simply be limited by another factor. Production however cannot be increased to an unlimited degree simply by adding what is deficient. It is true: a pumpkin can only reach a certain maximum size and no more. ‘This concept is incorporated in Liebig's law of the minimum, which postulates that the growth of a plant is dependent on those nutrients that ‘occur in minimum amounts. It would similarly apply to all organisms and environmental factors. ‘The identification and supplementation of short- ages of micro-nutrients in plants and animals has enabled large areas of land to be changed into more productive and economically viable agricultural land. In some cases factor compensation may take place. This happens when ideal conditions in terms of one factor compensate for conditions outside the normal tolerance threshold of another factor. Normally crayfish can withstand relatively high temperatures; if the salt concen- tration is below 14 parts per thousand and the ‘oxygen content is below 3mg/|. If these values are exceeded then they can only survive at a lower temperature of approximately 28°C. Chapter 3: The Dynamics of Ecosystem Functioning 31‘Average annval rantal (men) (Source: Css, 1950:47) Tolerance spectrum of biomes Biological modification of environmental condi- tions is also found in Nature. To a certain extent plants and animals are capable of changing some environmental conditions. During winter, temper- atures within boreal pine forests can be at least 2 or 3 degrees higher than on the surrounding snow covered grassiands. During summer the opposite can happen. To a certain extent plants can "create" and maintain their own micro- climate. ‘Trees radiate heat and slow down air movement so that the effect of wind and exposure is lim: ited. In misty areas condensation and dripping water from trees can increase the amount of precipitation reaching the ground surface. The crowns of trees can also influence the speed at which precipitation reaches the ground. Plant cover also influences soil temperature and evaporation. A good plant cover prevents direct rays of the sun from reaching the soil, while evaporation from the surface of the soil also decreases (see Gaia-hypothesis par. 11.2.2). Before any fluctuation can cause a change in the distribution of organisms, the extent of the change must exceed the ecological threshold (or tolerance level) of the organism in order to overcome internal resistance to change. This internal resistance results partly from the ability of organisms to create their own micro-climate. The vegetation of the rain forests of the Amazon Basin can only be maintained if the annual rain- fall exceeds 2 000mm. Only about 1 000mm of this rainfall originates over the Atlantic Ocean, the rest is provided by evapo-transpiration from the plant cover itself and from the soil. When the plant cover is destroyed to such an extent that the contribution from evapo-transpiration falls ‘average anew atl (re) (Aeptes rom: Cross, 1990) Figure 3.2(b): Tolerance spectrum of a temperate deciduous forest in terms of moisture and temperature below 1 000mm, positive feedback comes into operation. For all practical purposes it will become impossible to prevent the further decline of the forests or to even rehabilitate them. Similar relationships exist between organisms and the soil. Through succession, Nature is able to restore damage to an ecosystem, but only if a reservoir of the original climax community is maintained (par. 3.10). In the absence of a reser- voir, or when a number of species has become extinct, strictly speaking, a new climax will develop. It goes without saying that the larger the damage, the smaller or more isolated the reservoir, the longer it will take to reach a climax again. Pioneer communities on damaged areas can change the micro-climate and the soil in such a way that the reintroduction of the climax species can take place more easily. ‘The productivity, number, and distribution of plants and animals are controlled by five main ‘groups of factors: 1. Climatological factors, for example, solar radiation, temperature, humidity, precipita- tion, wind and exposure, 2. Edaphic factors, for example, physical characteristics of the soil, water relation- ships, soil chemistry and nutrient status. 3. Biotic factors, for example, the effect of grazing, fertilisation and trampling by animais; burning, deforestation, cropping or transport by humans. ‘4. Geological and evolutionary factors, for example, global tectonic movement and continental drift, long term climatic change. 32 cEcoplan Environmental Management5, Internal biological factors, for example, species strategies and competitive mecha- nisms, succession, health, age and geneti- cally based variations in tolerance or pro- ductivity. Some of these factors determine the spatial distribution of plants on the surface of the Earth. Species with a large tolerance range with respect to climatological, edaphic or biotic factors and which are thus widely distributed are known as eurytopic. Species with a narrow tolerance range only occur in localised areas with specifi cally favourable conditions and are considered to 3.1.4 Trigger factors ‘The principle that any human action results In a chain reaction causes human impact to have more complications than is usually anticipated. Bringing water into drought stricken areas for irrigation will decrease loss of life among livestock, but it also leads to over grazing, over stocking and erosion, which in turn results in the loss of fertile top soil. This then leads to further weakening of pasture (grazing), bush encroachment, etc. Changing a factor within a system will obviously have a much more complicated impact than the removal of an element from a static structure. The addition of water to the petrol of a car will eventually lead ee 3.1 be stenotopic in their distribution. It should be noted that eurytopic does not necessarily always relate to tolerance. It also relates to geographic distribution. Geographic boundaries are important limiting factors. Certain fresh water fish, have, for example, a very limited distribution because under normal circumstances they cannot move from one pool or river to another - if they are, however, moved artificially, they may flourish in their new surrounding, very often because their ratural enemies are no longer present. Certain eurytrophic species may therefore appear in practice to be stenotopic in their distribution. to disastrous side effects but there will be less significant effects if one should remove a brick from a wall. Adding excessive nutrients to a water body will lead to eutrophication (enrichment) of the water, stimulating the growth of algae and water plants to the extent that oxygen levels will deteriorate which will affect aquatic life and ultimately human welfare. Industrialization has similar vicious side-effects which place great stress on the environment as a resource. The principle of "trigger factors” therefore caution us to be careful of any changes to the existing environment. The cat population on Marion Island: the problem of invaders The effect of external influences on sensitive ecosystems is well illustrated by the case of Marion Island. In 1949 five cats (Felis catus) were broughit to the weather station as pets. Within 28 years the cat population has increased to 3 400. This increase was due to the absence of natural enemies and the superabundance of food in the form of birds that were completely oblivious as to the dangers of land-based predators. The fauna and flora of Marion island, being of recent volcanic origin, has developed through trans-marine migrations. It is therefore Chapter 3: The Dynamics of Ecosystem Functioning 33a relatively poor and specialized biota with no terrestrial predator species. The island thus became a haven for seabirds and seals, The food chain is simplistic but highly sensitive towards invaders. The cats stepped into this "utopia’ and in one year killed approximately 450 000 birds, especially the Pelecanoides urinatrix. This has had an influence on the whole food cycle on the island, The Chatharacta antarctica, which is one of the few predator bird species, was severely inhibited in its feeding habits; turning to penguins as food source. The removal of so many birds also had a detrimental effect on the amount of organic fertilizers (qwano) left by the birds and this influenced the’ vegetation negatively by reducing e.g. concentrations of nitrogen drastically. Apart of this, less birds also have had a whole series of impacts on micro- organisms, macro-invertebrates such as earthworms, snails and insects and through this, the soil and vegetation was negatively influenced. The methods of getting rid of the cats by introducing a virus and later hunting the rest, is discussed elsewhere (Gildenhuys & Hugo; 1991). Of interest is that even the secondary activities associated with the eliminating of the cats, have had a trigger effect on the ecosystem. The footpaths trodden by hunters gave tise to compaction and invasion of exotic grasses such as Poa pratensis. The trigger effect of this on the fauna and flora cannot be curtailed. Techniques of measuring environmental impacts are being discussed in chapter 10. 3.1.5 Holocenotic principle ‘The environment is not a simplistic system in which every element can be studied on its own and where cause and effect can be calculated like a chemical reaction. The ecosystem is too intricate and complex for this. Our knowledge of the Earth is too inadequate to carry on the process of large-scale environmental change and yet still avoid a world-wide catastrophe. This question will have to be approached from a holistic point of view. It is ironically a little-known fact in South Africa that Gen. J.C. Smuts was the initial proponent of the theory of holism. Through his epic work Holism and Evolution, 1926, he established himself as one of the important founders of ecology. Perhaps his greatest contribution lies in the fact that he saw how humankind was equally integrated into the total environment with all plants and animals. Just how great was the contribution of Smuts to science? Perhaps being the recipient of ‘twenty seven honorary doctorates says some- thing. Einstein is sald to have reported in 1936 that he (Smuts) was one of eleven people who truly understood his theories of relativity. In summary, holism teaches us that every- thing is attached to everything else in a func- tional, dynamic way. These intricate relation- ships can only be studied by viewing the Earth as a system, If humans are ever to succeed in thelr quest for sustainable living, then there needs to be a thorough understanding of this, system, and most importantly, of the role humans play as part of this system (see chapter 5 and 9) 34 © Ecopian Environmental ManagementA sicurr 31 Holism is best seen as the opposite of reductionism, or the "analytical method” which underlies modern science. The holistic approach consists of studying complete living systems, rather than the different components in isolation "in controlled laboratory conditions" (as modern science does). This is justified on the grounds that natural systems are more than the sum of their constituent parts. By studying the parts separately, reductionist science fails to take into account the way in which these components are organized. Holism ‘To insist on trying to understand biological and cultural phenomena in terms of physics or molecular biology (which is still the fashion in scientific circles), is thus considered by holists to be a vain pursuit. It does not enable scientists to understand the effects of the changes they are bringing about to our health, our society and to the biosphere, whose true nature can only be understood in the light of a holistic view of the world. (Goldsmith & Hildyard, 1990:92) The functioning 32? of environmental systems ‘n previous chapters mention was made of ‘concepts such as photosynthesis, respiration, producer and decay organisms etc, These concepts and their exact position and function in the environment as a system will now be discussed in more detail. 3.2.1 Green plants have the ability to produce complex ‘organic compounds from simple elements derived from the soil, water and air. Not only are plants dependent on this process, but every other form of life on Earth as well, including humans. In its simplest form this process comprises the production of glucose from carbon dioxide and water in the presence of light. This process known as photosynthesis (photo = light; synthesis = join). In the above-mentioned reaction 2 770 kilojoules (Kd) of solar energy in the form of light is needed to produce one Food chains TCOME 32 You should understand food chains and energy flow through the ecosys- tem: food chains being circular and energy, one directional. molecule of glucose and six molecules of oxygen (see Fig. 3.3). This energy is available to the plant itself as well as to animals feeding on the plant. Photosynthesis takes place in the chloroplast, Where a pigment called chlorophyll is present. In the presence of chlorophyll water and carbon dioxide combine to form glucose. By means of further processes and in the presence of a large variety of enzymes, other nutrients such as proteins, fats, starch etc, are formed. Chapter 3: The Dynamics of Ecosystem Functioning 35ISUN | ——— (Chloroplast te (1:0) ane carton coe (©or) so apt and goose caron dioxide | | Stucose (Cot204) Is synthesised CO, CoH 20e CO; + GH;O + light + CeHy20y + 602 Figure 3.3: A model of photosynthesis INSIGHT : i7 Photosynthesis When light strikes a chlorophyll molecule, it is absorbed and its energy is transferred to an electron of the chlorophyll molecule, now raised to a higher energy state. Once raised to this higher energy state, the electron is passed along through a series of chain reactions. This results in a synthesis of adenosine triphosphate (ATP) from adenosine diphosphate (ADP) by forcing a third phosphate group onto ADP. At the end of this reaction the electron, drained of its excess energy, is passed to a compound called trophosphopyridine nucleotide (TPN). Acquiring this electron gives TPN a strong affinity for hydrogen irons, available from the ironised form of water (H'+ OH). The energized TPN apparently pulls two hydrogen ions from the water molecule, splitting the water and forming TPNH,, and retains the electron. The remaining OH ions form water, release some molecular oxygen, and supply the electron to replace the one lost by the chlorophyll molecule. The electron is then passed along a series of acceptor molecules until it arrives at the chlorophyll molecule. Further reactions involve the synthesis of carbohydrate from CO; in another series of reactions that incorporates the energy as well as.the hydrogen and CO, into carbohydrates. (Smith, 1990:31-36) 36 otcopian Environmental ManagementBecause green plants are able to manufacture their own nutrients and are not dependent on other organisms for food they are called auto~ trophic or self feeders. They are therefore the primary producers of nutrients in the eco- system. All other organisms are either directly or indirectly dependent on the autotrophs for their nutrient requirements and are known as hetero- trophs (the food chain has been illustrated in Fig. 1.4). Heterotrophs can be divided into a number of groups, namely: @ _ herbivores or plant eating animals (browsers and grazers); ® carnivores or meat eating animals; @ _ saprophytes or plants that live on dead organic material; @ — saprozoa or animals which feed directly on dead organic material; @ omnivores which eat both plants and meat (or insects etc.); and @ parasites which are both plants or animals which live on other plants and animals. In contrast to carnivores, parasites do not neces- sarily cause the death of the host organism. Parasites can live on the tissue of the host organ- ism, for example, mosquitoes that are blood- suckers, or they can live in the digestive tracts of their hosts without causing any damage, for example certain roundworms. They only rob a portion of the food of the host organism. Itis therefore obvious that a large number of organisms are dependent on other organisms for their food requirements. A herbivore such as an antelope feeds on grass or leaves, but can become the prey of another animal such as a lion. This mutual connection is known as a food chain (Fig. 1.4), while every link in the chain is known as a trophic level. Plants form the first trophic level (or T1), the antelope the second trophic level (or T2) and the lion the third trophic level (or 73). It is important to note that the division into trophic levels is a classification based on function, and not on species. One and the same species, for example humans, can occupy different trophic levels. Omnivores such as baboons can, for example, occupy levels T2, T3 and T4 while insect eating plants can occupy levels T1 and T3. It is possible to distinguish two main types of food chains, namely, a grazing food chain and a detritus food chain. In the first instance, green plant material, seeds and fruit (as well as dry grass and leaves) are eaten by herbivores which in turn may become the prey of a variety of carnivores. In the second instance, dead ‘organic material forms the food of the decom- posers. These decomposers (the saphrotrophic level) are also subject to a large number of predators that feed on them. It is again possible to distinguish two types of decomposers, namely the macro-decomposers and the micro-decomposers. Macro-decom- posers include organisms such as earthworms and wood lice, while a number of organisms that live on the excretions of other animals are called coprophagic, for example, dung beeties. A large number of fly species complete part of their life yCles in rotting organic material of plant or animal origin. ‘The micro-decomposers consist of a large variety of bacteria, fungi and other single celled micro- organisms. Fig. 3.4 Illustrates a terrestrial grazing food chain, Eragy so poe Brau aa Bes f ee HOW iene pensoaryc) ae, | (Comptie rom wnt et 19843154) Figure 3.4: Terrestrial food chain and energy flow Chapter 3: The Dynamics of Ecosystem Functioning 373.2.2 Food networks Food chains very seldom occur in isolation, but rather join up to form an interconnecting food network or food web. A simple food web can be as follows: a partridge eats grass seeds and grass sprouts as well as a caterpillar that was also eating the same grass sprout. The partridge also eats a few maggots that are found in the carcass of an animal that was caught by a lion. ‘The partridge can become the prey of a jackal, which in turn becomes the prey of a large bird of prey. However, the large bird of prey dies as a result of an infection and then becomes the source of food for a large number of organisms such as flies, beetles and bacteria, and over time, it is reduced to the original inorganic matter that forms the basic material of living organisms. Fig, 3.5 is an actual representation of a food web found on Marion Island and shows the effect of two exotic species in the food web. For a discussion of this food web consult Gildenhuys ‘and Hugo, 1992. 3 Cycles in the 1e ecosphere can be viewed as one large cycle: birth, (germination), growth, reproduction, death and decomposition out of which new life spring up again. It is not only eneray and nutrients that flow through the ecosystem from trophic level to trophic level. All matter is involved in a continuous process of movement and change. Three main types of other physical cycles can be distinguished, namely the hydrological cycle, a gaseous cycle and the sedimentary cycle. The Tatter two cycles involve both living organisms as ‘well as chemical processes and are known as biogeochemical cycles. Underlying each cycle Is a reservoir where elements or compounds are stored for various periods of time before they once again become part of the cycle. In the case of the hydrological cycle the ocean is the main reservoir, the atmosphere is in most cases the reservoir for the gaseous cycle while the Earth forms the reservoir for the sedimentary cycle. Within each cyde a number of smaller cycles can also be (Giienhuye @ Hugo, 1992 Figure 3.5: Food chain on Marion Island in terms of exotic cat and mouse populations ecosystem TCOME 33 You have to understand and be able to discuss the continuous movement of elements through the system to ensure that ecosystem health and stability is retained. Material lows from the living to the non-living and back to the living parts of the ecosystem in a perpetual cycle. In these cycles plants and animals acquire some of these elements (the macro- nutrients) in relatively large quantities. Others (the trace elements, or micronutrients) are needed in lesser, often only minute, quantities. Examples of cycles will be discussed briefly. Environmental Management3.3.1 The hydrological cycle The hydrological cycle is the best-known and most noticeable cycle in Nature. In its simplest form it consists of water that evaporates from the surface of the ocean and this moisture is carried to the continents where it condenses, forms clouds, and can later fall as some form of precipitation to the Earth. Part of this water sinks into the ground whilst about 70% again reaches the atmosphere as a result of evapora- tion and transpiration. The rest eventually reaches the sea as run-off via rivers and the whole process is repeated. The hydrological cycle can be viewed as a giant distillation process by means of which fresh water is made available to terrestrial ecosystems. A simplified form of the hydrological cycle is shown in Fig. 3.6 In spite of this apparent simplicity the hydro- logical cycle is one of the most important nutrient cycles (food cycles), and in some respects one of the most complex in the eco- system. The hydrological cycle is not uniform over the surface of the Earth, but changes according the various climatic zones. It also exhibits seasonal as well as long-term variations. Forms of life within an ecosystem are adapted to the specific nature of the hydrological cycle on . Some plants can withstand long periods of 3.3.2 Gaseous cycle drought, whereas others are not drought resis- tant (Principle of Limiting Factors). Some plants are dependent on ground water, while others need open water surfaces. Plants are dependent on dissolved inorganic salts present in water, either in shallow surface water, ground water or in underground reservoirs. Ground water content is dependent on factors such as the rate of infiltration, gradient, porosity of the soil, texture and rock type (Holocenotic Principle). kB @ The Carbon Cycle | FOSSIL FuELs ——) Figure 3.7: A simplified carbon cycle Carbon, together with hydrogen, forms the basic building block of all organic compounds, and therefore of all life on Earth. It is also the main ‘component of fossil fuels and, together with calcium and oxygen, forms the basis of all carbonate rocks. In most cases carbon has a very short cycle or circulation period from its two most important reservoirs, namely the atmosphere and the sea, to the plant and animal world. The atmosphere contains carbon in the form of carbon dioxide (CO:) at a concentration of approximately 0.03-0.04%. The ocean contains 30-50 times more CO: than the atmosphere and serves as a buffer or "sink" where surplus CO: is absorbed and subsequently the atmospheric CO: |s maintained at reasonably constant levels. Carbon dioxide is removed from the atmospheric and oceanic reservoirs by green plants through the process of photosynthesis. Its then stored in their tissues in the form of ‘organic compounds. Carbon dioxide is returned Chapter 3: The Dynamics of Ecosystem Functioning 39to the reservoirs through the process of respi ration by piants and animals (see par. 3.4), by burning organic material such as grass and wood, and more recently by burning fossil fuels, as well as the decomposer organisms that finally break down organic material into various components. Carbon also moves through the ecosystem from the primary producers, namely plants, to the herbivores, and on to the carnivores. At death, decomposer organisms break tissue down to carbon dioxide and inorganic compounds (see also par. 3.2). The carbon cycle can be viewed as a nearly perfect cycle because under natural conditions almost as much carbon is set free through the process of respiration as is bound by photosynthesis. However, due to industrialization, the burning of fossil fuels releases enormous amounts of carbon dioxide in the atmosphere. This causes the so-called "Greenhouse" effect that in tun is aggravated by Because oxygen is so reactive, it's cydiing is quite complex. As a constituent of carbon dioxide It Grculates freely throughout the biosphere, Some carbon dioxide combines with calcium to form carbonates. Oxygen combines with nitrogen com- pounds to form nitrates, with iron to form ferric, ‘oxides, and with many other minerals to form various other oxides. In these states oxygen is temporarily withdrawn from circulation. In photo- synthesis oxygen is split from the water molecule, During respiration by plants and animals, oxygen ‘combines with hydrogen to form water. Part of the atmospheric oxygen that reaches the higher levels of the troposphere is changed to ozone (0) by high-energy ultraviolet radiation. The nitrogen cycle is summarized by means of a diagram, Fig. 3.8. For a description, see Smith, 1990:258. Due to the importance of micro-organisms in the system, the role of nitrogen should not be underestimated. deforestation. These aspects will be dealt with in subsequent chapters. In the case of plants, the re~ circulation period can be less than 24 hours. In the tong term carbon can be fixed in the exoskeletons of certain organisms in the form of carbonates, for example coral in the sea, and can only be retumed to the atmosphere over long periods of time by chemical processes. A simplified carbon cycle is illustrated in Fig, 3.7. @ Oxygen and nitrogen cycles The oxygen cycle is another example of a well-known gaseous cycle. Oxygen, the by-product of photosynthesis, is involved in the oxidation of carbohydrates with the release of energy, carbon dioxide and water. Oxygen is very active | chemically. It can combine with a wide range of chemicals in the Earth's crust and is able to react spontaneously with ‘organic compounds. De vt ‘Vosan aston Saoera se a SS /: we Prien ss ad /) Sel cecomposing) Bacna nieaior as (Aeapted tom: Smith, 1977:84) NOs by rate bales > Figure 3.8: Nitrogen cycle 40 Econ Environmental Management3.3.3 Sedimentary cycles Minerals required by living organisms are initially obtained from inorganic sources. Available minerals ‘cour as salts dissolved in ground water or open water on land, All sedimentary cycles consist of ‘two phases: the salt solution and the rock phase. Mineral salts come directly from the weathering Of rocks from where it enters the water cycle and eventually reaches the sea where it remains indef- initely. Other salts accumulate in the Earth's crust through sedimentation in the form of silts and limestone. After weathering they are retumed to the cycle. @ The phosphorus cycle ‘The phosphorus cycle is an example of a typical sedimentary cyde. In contrast to the hydrological and gaseous cycles the atmosphere plays no role in the phosphorus cycle. Phosphorus is an impor- tant component of nucleic acids in all cells, and as part of adenosine tri-phosphate (ATP), it is the form in which energy is used by all forms of life. Phosphate plays an especially important role in animals because it constitutes about 50% of bone. ‘The movement of phosphorus through the eco- system takes place as follows. Weathering breaks down rocks containing phosphates and the phos- phorus is set free into the soll It is then absorbed by the roots of plants and incorporated into the cells. Animals then obtain their phosphorus by feeding on plants. In some cases, especially in hoofed animals, (or ungulata) supplementary phosphate is obtained by means of "salt licks", Camivores or meat eaters ingest the bones of their prey to satisfy their needs. The terrestrial eco- 3.3.4 Circulation periods These are the estimated times it takes for the available supply of a certain element to circulate once through the ecosystem. It must be distin- guished from the time It takes a single molecule to complete a cycle. ‘Some molecules will complete the cycle many times during a process, while others will not even complete the cycle once. In the case of carbon it is estimated that an amount of carbon equal to the total world supply will move through the ecosystem once every 300 years. A number of circulation periods are indicated in Fig. 3.9, but the data in this figure is highly speculative, system loses tonnes of phosphates to the oceans annually through erosion. Dissolved phosphorous, salts are absorbed by phytoplankton and made available to larger animals by means of the marine food chains. A part of this phosphorous is eventually returned to the land in the form of excreta (guano) of fish eating birds but this is limited to the coastal regions, Moder technology and associated human activities have caused a tremendous disturbance in the natural phosphorous cycle. Large amounts of phos- phate containing rocks are mined and by treating these rocks with strong acids such as sulphuric acid, the relatively insoluble di- and tr-calcium phosphates are changed into highly soluble mono- calcium phosphates. These are used on a large scale as synthetic fertilizers to obtain higher agrl- cultural yields. Eventually, by means of the sewer- age systems of large cities, these phosphates end up in rivers, dams, lakes and the sea. Over time part of these phosphates combine with other dissolved salts and sink as sediments onto the sea bed where, for all practical purposes, its lost to ‘ordinary ecosystems. ‘A further source of phosphate in ecosystems is the vast amount of detergents agents used by people in various washing processes. This phosphate, reaches water sources where it causes eutrophi cation, with resultant algae blooms. 2 Other sedimentary cycles Calcium, potassium, sulphur, magnesium and other tycles can also be distinguished (see Smith, 1990). 0 coe (Erion ewe Figure 3.9: Circulation periods Chapter 3: The Dynamics of Ecosystem Functioning 414 Energy flow and food pyramids Tis Scene La of Termodynamis tach us that energy cannot be used again because it loses its quality when changed from one form to the other - the principle of entropy. With every transfer of nutrients from one trophic level to another a large amount of energy is thus lost; mainly in the form of heat during the process of respiration. In this case high quality chemical energy is converted to low quality heat energy. Fig. 3.10 fsa diagrammatic representation of the flow of energy through an ecosystem. Respiration is the opposite of photosynthesis and comprises the oxidation (burning) of nutrients with the release of energy. The organism can then use this energy for the maintenance of its life processes, ‘The following reaction represents the process of respiration: CéHHnOs + 60: = 6CO: + 6H. + energy Bp = Gross primary production Np = Nett primary production Secondary production (at trophic levels 2 and 3) E = Energy received sp = 7 z f 4 i ‘ouToTROPHIC | | wevERo. eae | TROPHIC 509 Bp Nee = ‘Sp2. = z at ‘SUNLIGHT 5 ge | ) | | L ° Je ; i 3 Siday: - 124 62 6.2 V2 Se (Be)_(No} (Sp2) (8e3) SO ight Gh respiration Oy unused energy Wr heat Figure 3.10: Energy flow through the ecosystem Itis clear that there must be more, (or a greater biomass) of primary producers present in an Vacs ‘ecosystem than herbivores, and there must also PSP _ [TS retary camvores 5 be more herbivores than carnivores. In aquatic = secondary carnivores ecosystems there are varying levels of carnivores. Ff T2 Herbivores a reserved for the decomposer organisms and top | 3 4! ouro- carnivores. There should be fewer camivores in T4 | TH Primary prowess TROP compared to T3, etc. This decrease in the number of organisms from the T1 level to the TS level can be illustrated diagrammatically in the form of a Figure 3.14: Food chain pyramid 42 eecoplan Environmental Managementfood pyramid (Fig. 3.11). It is, however, not always correct to take only the numbers present in each level into account. In the case of insects, a large number can live on a single tree, and this can com- pletely change the form of the pyramid. It would be more correct to compare the mass present in trophic levels with each other. This is an example of 2 biomass pyramid. Fig. 3.12 represents a number of ecological pyramid: Pyramid a indicates a grassland with herbivores and carnivores, pyramids (b) and (d) show a forest with insects as primary consumers, while pyramid (c) represents a biomass pyramid. A biomass pyramid may also be “inverted” as in the case of whales living on plankton. Pyramids (@), (b) and (4) are based on numbers. Hyper- parasites (Fig. 3.12(d)) are found on other parasites. The only pyramids that always have a broad base are energy pyramids. The decrease of energy with each increasing level has enormous implications for the increas- ing world population. Many more people can be maintained on a certain area if humans only occupy the level of herbivores on 72, rather than that of primary carnivores on level T3. It would, therefore, be more advantageous if the yield from a piece of land, for example @ maize field, were for direct human consump-tion, rather than to first feed it to livestock such as cattle or pigs and then use the animals as food (meat) or food products (milk and eggs). Grassland ecosystem with mammals, birds, etc. as primary consumers fe seed NUMBERS PYRAMID Testiary camivores, Secondary carnivores Primary carnivores © aaa eata! sertores a PIETY "Producer erie BIOMASS PYRAMID (also eneray pyramid) Forest ecosystem with insect as primary consumers (b) 3] Tertiary combores @ toy epgereions Comoe EEG ie mente 2S] Pinar orate z NUMBERS PYRAMID NUMBERS PYRAMID Figure 3.12: Ecological pyramids Protein derived from plants is unfortunately not sufficient to satisfy the total nutritional requirement of humans. Plant protein is not 2 “complete” source of protein because certain essential amino acids are not present and humans cannot synthesise these amino acids on their own, Many animals especially herbi- vores, are however, capable of synthesizing these amino acids, and by using animal products humans can satisfy all their protein requirements (par. 5.1.3). The extensive grazing areas of the world cannot be used for the successful production of crops such as maize, beans, rice etc. These areas can, however, be used for livestock farming and thelr products used by humans. The yields from areas where crop farming is possible must rather be directly used by people and only supplemented by products from the extensive grazing areas. In this way the food requirements for the present world population can be met with ease. Chapter 3: The Dynamics of Ecosystem Functioning 435 Productivity and biomass ‘t has been shown that in Nature, there Is never stagnation. Everything is always in a 3.5.1 Primary production process of change. Growth and decay are constant, ‘The flow of energy through the community starts with the fixation of sunlight by plants, a process that in itself demands the expenditure of energy. Plants start to grow by living on the food stored in the seed until its production machinery is working. ‘Once started, the green plant begins to accumulate energy. Energy accumulated by plants is called production oj, more specifically, primary pro- duction, since itis the first and basic form of ‘energy storage in the form of complex organic compounds in the tissues of the plant. The rate at which energy accumulates is known as primary productivity. All of the sun's energy that is assimilated - that Is, total photosynthesis - is gross primary production. Since plants, lke other organisms, must overcome the tendency of energy to disperse, free energy (that is available to do work) must be expended for production as well as, for reproduction and maintenance. The energy required for this is provided by the reverse of the photosynthetic process: respiration. The energy remaining after respiration and stored as organic matter is net primary production, or plant growth. Therefore the net production for a single plant or a plant community can be described by the following formula: Net primary production = gross primary production ~ autotrophic respiration ‘The basic unit of energy used in ecological work is the calorie, usually expressed as kilocalories per square meter per year (Kcl/m'/year). However, 3.5.2 Secondary production production may also be expressed as dry organic matter in grams per square meter per year (a/mm'/year). Net primary production accumulates over time as plant biomass. Part of this accumulation is tured over seasonally through decomposition. Part is retained over a longer period as living material. ‘The amount of this accumulated organic matter found on a given area at a given time is the standing biomass. Biomass is usually expressed as grams of organic matter per square meter (g/m), calories per square meter (cal/m?), or some other appropriate measure per unit of area, Thus biomass differs from production, which is the rate at which organic matter is created by photo- synthesis. The biomass present at any given time is not the same as productivity. Biomass, both living and dead organic matter, varies seasonally and even daily. Biomass is thus the mass (weight) of living organisms per surface area or per volume Unit, The total number of organisms within a certain area is known as the biotic community and their mass is the total biomass of the com- munity. Biomass can also be expressed in terms of a specific species, for example the number of elephants in the Addo Elephant National Park, or red grass (Themeda triandra) in a certain grazing area. It indicates the mass of a certain species that is available in the ecosystem under discussion. The portion of the biomass, which is available as food for subsequent trophic levels, would be referred to as the standing crop. ‘The energy derived from the gross primary pro- ductivity can be accumulated in the ecosystem and serves as a basic source of food for primary con sumers. The amount of energy accumulated by the primary consumers, or the T2-organisms, is known as the secondary productivity of the ecosystem. Once consumed, a considerable portion of the plant material, again depending on the kind of plant involved and the digestive efficiency of the herbivore, may pass through the animal's body undigested. A grasshopper assimilates only about 30 percent of the grass it consumes, leaving 70, percent as waste. Mice, on the other hand, assimilate about 85 to 90 percent of what they consume. 44 © Ecopion Environmental ManagementEnergy, once consumed, is either diverted to main- tenance, growth, and reproduction or is passed from the body as faeces and urine (see Fig. 3.4). Depending on the nature of the organism, the loss of energy can be variable and often quite high. ‘Another portion is lost as fermentation gases. OF the energy left after losses through faeces, urine ‘and gases, part Is utilized as “heat increment,” ‘which is the heat required for metabolism above that requited for basal or resting metabolism. Population numbers 36 and carrying capacity Il organisms have the ability to produce more than two offspring so that at the end of the life span of one set of parents, there should be more offspring than the original set of parents. Animals with a slow reproduction rate, like elephants, can produce one offspring every four years for approximately 40 years. That is approximately 10 offspring per set of parents. If itis accepted that elephants reach reproductive stage at 16 years of age, it means that the elephant population can increase by a factor of five every 56 years. Birds can produce between one (albatross) and 40 (quail) offspring per year; while amphibians can produce a few hundred, fish a few thousand and insects a few million offspring per year. Plants can produce between ‘two and a few million seeds per generation. As a result, in theory, all populations will increase. This is, however, not the case. In a stable eco- system populations tend to remain constant. ‘The population of any species at any stage is the difference between natality plus immigration and mortality plus emigration. Natality is the natural growth owing to birth, hatching or germination while mortality represents the natural death rate. Because environmental conditions are seldom ideal, populations never reproduce at the theoretic potential or biotic potential. Mortality in the immature population is usually very high. Competition and preda- tion are the most important factors that prevent all offspring from reaching the reproductive stage. Ina stable ecosystem, on average, only two offspring, Irrespective of the number produced, eventually reach maturity to replace their parents. Otherwise population numbers would change over time. The growth of a population can be represented graphically by means of two graphs, namely the S-shaped or sigmoidal curve and the J- shaped curve (Fig. 3.13). The S-shaped curve is TCOME 3.4 You need to be able to describe how population numbers are being kept in check by Nature. i canying capacty POPULATION INCREASE Figure 3.13: Growth curves usually found in populations with a long life span and a slow reproduction rate such as large mammals. The J-shaped curve is more often found among species with @ short life span and a high biotic potential, for example, insects. In the S-shaped curve there is initially a slow popu- lation increase but the rate increases gradually until the carrying capacity of the environment Is approached, after which point the population numbers will stabilize and a state of balanced equilibrium is arrived at. Unless changes in the environment occur, the population numbers will remain fairly constant over time. In the J-shaped curve there is initially a tremendous increase in the population so that the carrying capacity of the environment is quickly exceeded and a catastrophic decrease in population numbers Chapter 3: The Dynamics of Ecosystem Functioning 45follows. The equilibrium is of a more dynar nature, In the insect world such a situation often occurs at the end of the season, but some individuals may survive unfavourable conditions in the form of eggs, larvae and pupae. The periodic suicidal migrations of lemmings, ‘a guinea pig-like animal of Northern Europe, are another example. These animals have a very high potential and as a result their numbers increase at a very high rate during 5 to 7 seasons so that the carrying capacity of the environment is greatly exceeded, In order to overcome this, lemmings begin a migration process which can apparently not be stopped once it has begun. Large numbers become the prey of predators and birds of prey; some drown as they cross rivers while yet others fall off cliffs or may die of hardship during migration. ‘The survivors eventually reach the coast and apparently fearlessly rush into the sea and drown. Before humans destroyed the habitat of the lemmings this migration process reached spectacular proportions. A relatively small number of lemmings do not take part in the migration and serve as @ reservoir from which population numbers can again begin to increase Until the whole migration process repeats itself in a few years time. In this case the growth curve (Fig. 3.13) will decrease at an even sharper rate than indicated on the graph. ‘The carrying capacity (par. 3.11) of the envi- ronment or ecosystem Is those population numbers that can be maintained by the resour- ces of the ecosystem. If the carrying capacity of the ecosystem is exceeded, then physical and biological contro! mechanisms will be activated so that environmental resistance starts to build up and, as a result, population numbers will decrease or stabilize on a lower level (compare negative feedback, Fig. 1.3). Biological control mechanisms for population jcal control mechanisms for population num- bers include factors such as a shortage of food, especially during dry periods that eliminate the weak- 3.7.1 Predation numbers est individuals; other examples are a shortage of sheltes fire, ete. These control mechanisms cause complicated intra-species and interspecies interactions. Predation is the interaction whereby one orga- nism, the predator (e.g. lion), kills and devours another organism (buck), the prey. The typical predator is therefore larger than the prey, or occasionally two or more predators can work together to bring down the prey, for example wild dogs. With the exception of certain insect eating plants predation only occurs in the animal kingdom. Predators fulfil an important role in the com- munity. Weak, sick, maladjusted or injured animals are the first victims of the predators. Only the strongest survive and this ensures that ‘any defects present in the population are eliminated at a very early stage. In order to survive, specialized adaptations are required by both predator and prey. ‘Adaptations enabling escape from predators are known as survival strategies and include camouflage, and escape through speed, or reproduction strategies, such as a tremen- dously high biotic potential, for example aphids, Which reproduce at such a high rate that the predators are simply not capable of endangering population numbers significantly. ‘One of the most promising applications of pest control is the principle of biological control by making use of certain predators to control agricultural pests (see par. 5.3.2.2.1.2). 3.7.2 Cannibalism, cronism and prolicide Many examples exist in Nature where adult ani- ‘mals, without apparent reason, kill and eat their young, Although such behaviour in the animal kingdom is difficult to understand, itis certainly not unusual. This phenomenon is known as cannibalism and can sometimes be seen as 46 oF:coplan Environmental Managementpart of a feeding strategy. There are approxi- mately 138 different kinds of animals that are known to practice cannibalism in one form or another; these vary from single celled protozoa to spiders, insects and molluscs to fishes, birds and mammals. It is important not to view canni- balism as deviant behaviour, but rather as a reaction to a wide range of environmental stimuli, With the passage of time a number of additional concepts besides cannibalism have been developed. A prolicider (derived from the Latin word prolis = descendent) kills the young, generally not its own. It can happen, for exam ple, where one or more male lions take over another pride and then kill all the young. The females then immediately come into oestrus again and are fertilized by the new males in the pride. This phenomenon ensures that the descen- 3.7.3 Symbiosis or mutualism dants of the dominant males (with a strong gene ool) survive and reproduce. If the killed young are eaten It is called cronism, Fratricide occurs when siblings kill one another. If the animals that have been killed were then eaten, it would be called cannibalism. Cannibalism (and similar phenomena) are in some instances related to population density, and/or the availability of food. In some cases the behaviour of the victims may irritate the aggres- sor or the behaviour may simply be a release valve for stress under unfavourable conditions. Not all relationships between species are how- ever negative or restrictive. Species often interact ‘with each other in a positive, often stimulating way. AA situation in which a close and often permanent and obligatory contact exists between two species is called mutualism. Mutualism is often termed sym “ving together" and includes mutualism, commensalism, and parasitism. is. Actually symbiosis means, 3.7.4 Commensalism and amensalism Often a one-sided relationship between two species ‘exists, in which only one benefits and the other is neither benefited nor harmed, Such a relationship is called commensatism. Among the commensals are epiphytes such as orchids, These plants grow ‘on the branches of trees, where they are nearer to the source of light; drawing nourishment from the humid air. Examples are the “old man's beard’, Spanish moss and orchids in the forests where conditions are moist, These, and other epiphytes depend on the trees for support only - manufac- turing their own food by photosynthesis. Other examples of commensalism are: @ Fungi of the species Mykorrhisa, which pene trate the roots of their hosts, e.g. pine and ‘oak varieties, Short thick roots are produced, and the roots are covered with fungi hyphae ‘The fungi use the carbohydrates from the roots and in turn provide inorganic foodstuffs for the roots. ‘® Most members of the family Proteaceae have an oleaginous appendage known as the elaiosome. When the seed falls, termites are attracted by the scent. The termites carry the seeds over long distances to their nests, and eat the elaiosome, but not the seed. The seed is possibly also protected against fungal in- fection by secretions from the termites. The high level of formic acid protects the seeds against bacteria and possibly also suppresses germination. When the veld burns, the seeds are protected underground, where they germinate in the fertile soll of the termites heap. An Argentine species of ant is also established in the fynbos at present. ‘This invasive species eats the elaiosome on the surface where the seed is left. The seed is then exposed to fire, insects and rodents, which prejudices the seed's survival. In the animal kingdom there are many examples of ‘commensalism: The crocodiles that breed along the Nile lay their eggs in the sand on the riverbanks. Birds, lke the Plover ("kiewiet") (Vanellus) and “dikkoppe” (Burhinus) as well as turties prefer to nest alongside these crocodiles for protection against snakes and leguans who prey on their eggs. Duck-bamacles attach themselves to the backs of whales and to the shells of horseshoe crabs. ‘The sucking-fish (Remora) attaches itself to the abdomens of sharks. In this way, it obtains free transport and feeds on small portions of the shark's prey. In contrast to commensalism is amensalism, a situation in which one individual definitely inhibits Chapter 3: The Dynamics of Ecosystem Functioning 47the other while remaining unaffected itself. By so modifying the environment the organism improves its own chance of survival. Amensalism commonly 3.7.5 Parasitism involves some type of chemical interaction with other organisms. With parasitism one organism, the parasite lives either temporarily or permanently on or in another organism, the hast. It obtains its food from the host. It is only a short step from amensalism to parasitism. In parasitism two animals live together but one derives its nourishment at the expense of the other, The parasite, however, does not kill the host, because this could also cause its own death. Parasitism occurs in both the animal and plant kingdoms. Animals and plants can be parasites of other animals and plants. Parasites are smaller than their hosts. Sometimes parasites are only occasional visitors ‘with the main aim to obtain food, like mosquitoes, but they can also be permanently dependent on their host organism, for example dodder on lucerne. Ectoparasites occur on the outside of the host whereas endoparasites occur inside. Parasites can live on the tissue of the host, such as, bloodsuckers, the half digested food of the host, for example certain nematodes or roundworms or they can live on the secretions of their hosts, as pupae that live in the nasal cavities of sheep and some other herbivores do. In many cases they inrtate the mucus membranes of the host in order to increase secretion. Parasitism Is also characterized by special adapta- tions by both the host and the parasite, The host takes certain preventative measures in order to resist the parasite while the parasite takes measures to overcome the resistance of the host. ‘The most successful parasite is the one that causes the least damage to the host. Parasitism, Ike predatism is dependent on popu- lation density. The greater the host population the easier it is for the parasite to spread. The lower the resistance of the host, the more vulnerable itis to the parasite. When the carrying capacity of the environment is exceeded many weakened indivi- duals of a species may be present which can more easily be attacked by parasites. Parasites may weaken the organism still further and it may then become the prey of a predator. In this way para~ sitism may also be viewed as a population regulating mechanism. Very broadly speaking one can summarize the above ‘mentioned control mechanisms by saying that in rmutualism the reciprocal effect is beneficial; in com- ‘mensalism it is mostly neutral in parasitism slightly negative and amensalism represents a definite negative relationship. The more Form of symbiosis Species A | Species B | exact relationships is summarized by Mutualism + +. means of Table 3.1. The table shows, iti * ‘amongst others, that in competition rit pat : |) | Seiyeneee! Competition - fo. between two species. In parasitism a Amensalism : : parasite gains (s tse postvely affected) but has a (slightly) negative effect on the + == detriment 0 = no effect other (host) organism. In mutualism both (Source: McKinney & Schoch, 1998) 3.7.6 Suicide species gain by cooperation. ‘Summary of biological contro! mechanisms A very strange occurrence is the mass suicide that ‘occurs from time to time amongst certain popula tions. It was shown above that tens of thousands of Lemmings suddenly, without warming, would migrate to the sea and plunge over the ciffs into the ocean, A similar phenomenon is the voluntary ‘swimming to shore of whales and dolphins. At times they have been returned to the deep waters by concerned people, without success - only to ‘swim onto the beach again where they die. In in- stances these "suicides" were coupled with off- shore military exercises where it is thought that radar sounds might be affecting the sensitive direc- tion finding sensor mechanism of these animals. 48 oEcoplan Environmental Management8 Competition for resources he resources within a certain ecosystem are not unlimited, and as population numbers increase, competition for essential life sus- taining elements also increases, for example, food, shelter, nests and in the case of plants, space and sunlight. Two kinds of competition can be observed, namely intra-species com- petition where individuals of the same species compete, and inter-species competition, where different species compete with one another for the same resources. Intra-species 3.8.1 Competition is usually stronger than inter- species competition because different species very seldom or never occupy identical positions in the ecosystem. It is a recognized ecological principle that two species cannot occupy the same ecological niche that is they cannot share the same food, shelter and breeding space. In such a case competition between the two will take place until one Is excluded. Competition amongst animals Many animals are territorial by nature. Individuals demarcate a certain area for themselves and will then defend it against occupation by another individual by means of display or even fighting. The song of birds is one method used to demarcate a territory. Some animals will mark their territory by urinat- ing along the boundaries or by marking the boundaries with secretions from special glands. The inability of a male animal to demarcate 2 territory makes it almost impossible to attract a female and to reproduce. Some male animals will collect a harem of females that he defends against the attentions of other males: weaker males are again prevented from reproducing. A social hierarchy ("pecking order") is established between social animals, for example, flocks, herds, etc. Because of his dominant position in the group the leader intimidates the other group members to such an extent that their ability to reproduce decreases drastically. Inter-species competition usually leads to behaviour changes by one or both species. Small variations in food preferences, habits, time of activity, etc, can lead to a decrease in ‘competition, 3.8.2 Competition amongst plants Plants cannot change their location once they are established. The elimination of intra-species competition is therefore aimed at the distri- bution of seeds in such a way that no com- petition for space takes place. Where a dense stand of seedlings does occur, competition for food, water and especially sunlight will eliminate the weaker seedlings. Overshadowing is one of the most effective methods of competitive elimination, especially during ecological succession. Special adaptations by plants can eliminate inter-species competition to a great extent. During winter months the ground beneath deciduous forests is often covered with flow- ering plants that are capable of completing their life cycles in a short period of time. As soon as the trees begin to bud and sunlight is slowly eliminated, these plants enter a period of rest, either as seeds, bulbs or other underground structures, and only appear again when the leaves start falling from the trees during autumn. Climbing plants are examples of plants that are adapted to acquire sunlight, while others, like ferns, are adapted to low light intensity. Strictly speaking, these examples can be seen as niche- differentiation rather than species competition. Chapter 3: The Dynamics of Ecosystem Functioning 499 Ecological succession Succession ‘A well-known occurrence in South Africa that can be oftentimes seen from the road is the prolific growth of cosmos (kosmos) flowers in the road verges where the ground has been scraped clear of its natural vegetation. It also appears on newly ploughed lands. When these ploughed lands are left without planting crops on them, they are soon colonized by invaders such as khakibos and caster-oil plants (olieboom or Ricinus sp.). If left to mature, various types of grass such as bluegrass enters the scene until such time as the original veldt grass e.g, buffalo grass (buffelsgras) eventually starts to grow and replace the various seral stages with the ultimate climax vegetation type. These seral stages have little nutritional value for grazing, which is Nature's way of alleviating the area from further usage in order to restore the normal biota. ‘The profusion of pink, purple and white created by cosmos alongside the roads, is a well-known sight in South Africa. Less impressive, but as well known is the occurrence of khakibos on aban- doned fields. Wherever the natural vegetation has been disturbed, herbaceous plants that are notably different from the “normal” occur. If the original vegetation is forest, shrubs will soon follow the intial herbaceous plant growth, and eventually, after a number of years, trees will re- establish themselves to form the original forest. Thus over a period of years one community replaces another until a relatively stable forest finally occupies the area. ‘The changes involved in the return of the forest are not haphazard but orderly, and barring disturbance by humans or natural events, the reappearance of the forest is predictable. This orderly and progressive replacement of one community by another until a relatively stable community, (often called the climax), occupies the area is called ecological succession. The whole series of communities, from grass to shrub to forest, that terminate in a final stable com- munity Is called a sere, and each phase is a seral stage. Every seral stage represents a community, although temporary, with its own characteristics, which may remain for a very short time or for many years. When this process takes place along the shores of fresh water bodies, the stages are called hydroseres; along brackish water bodies the stages are called haloseres; on sand dunes itis called psammoseres; on ground, pedoseres and on rock surfaces lithoseres. In an abandoned cropland, succession obviously ‘occurs on one specific place with distinct boun- daries. In the case of a scree (talus) slope at the foot of e.g. the Magaliesberg escarpment, leading down into a valley, the different seral stages can be seen - the one "flowing into” the other. Near to the clifredge, where the debris consists of an abundance of large loosely compacted rocks with underdeveloped soils, a dominance of trees and large shrubs may be found. This vegetation diminishes down slope as the soll gets deeper and the soil horizons better developed, until, on the upper pediment (below the contact zone between plain and scree slope), short shrubs are steadily replaced by grass on the lower valley bottom. ‘The same principle of seral stages can be seen next to a river that is a dynamic changing geo- morphic unit; or a pond/viei that is gradually being filled up (see Smith 1990, p. 639), or sand dunes next to the coast (Kok, et al. 1987). 50 ocoplan Environmental Management3.9.1 Primary and secondary succession Primary succession is the establishment of pioneer species on or in a newiy formed habitat as happens, after a lava flow, or on a new island such 2s Surtsey Island in the northem Atlantic Ocean. Primary succession is normally (though not necessarily always) characterized by hardy species with a simple life cycle, a small number of species where one of, at the very most a few species, dominate the environment, a high reproduction rate, a small biomass and simple, short food chains, These plants change the environment by promoting the weathering of rocks, stabilizing the soil, adding of organic ‘material, which in turn increases its moisture retention capacity, and thus improves the nutrient status of the soil, Pioneer species establish themselves because ‘competition is usually absent. Later on plants that have traits enabling them to become estab- lished on previously modified environments replace them. ‘As succession progresses, the number of species present tends to increase and reaches a peak during the middle development phase, after which a slight decrease in species diversity can occur until the climax stage is reached. Secondary succession is the establishment of pioneer species in habitats that were previously populated. Examples of places where secondary succession occurs, are fallow lands, road verges ‘and cuttings, areas disturbed by fire, excessive erosion, overgrazing and excavations. The species most likely to colonize such places are mostly the so-called "weeds" and indigenous pioneer plants. Although virtually Indefinable, weeds usually have two common characteristics. They invade areas modified by human action and they are exotics, 3.9.2 The climax stage not native to the region. Once native species invade the area, the "weeds" eventually disap- pear, as they cannot maintain thelr dominance very long, Typical plants that emerge as pioneer species are "knapsekérel" (Bidens hippinata), khaki weed (Tagetes minuta), the castor oll plant (olieboom") (Ricinus communis), couch grass ("blou kweek") (Cynodon dactylon), bugweed (Solanum) (“luisboom") and cosmos (Bidens formosa). Indigenous plants will of course replace many of these species in follow-up seral stages, but aggressive invaders utilize such disturbances to establish themselves permanently lke the ‘Australian Acacias (Port Jackson) of the Cape Flats. Because pioneer plants normally are annuals and have to renew their photosynthetic structure each year, they eventually lose their dominance and are suppressed by the taller more vigorous growth of perennial plants at later stages of development. ‘These gradually assume dominance and are better able to exploit the site once the disturbing factors are removed ‘Secondary succession takes place at a faster rate than primary succession and is often charac- terised by a larger number of pioneer species. Its composition may reflect species from various seral stages. If changes to the environment have been initiated by external influences such as the draining of a ‘wetland, the change that will take place is called allogenic succession. When the plant com- munity itself alters the habitat, the process is referred to as autogenic succession. In this case one stage of plant development modifies the habitat to such an extent that itis replaced by another stage. Thus succession, especially primary succes sion, starts with the colonization of an area by Ploneer species. These plants are able to grow ‘on a substrate low in nutrients and organic matter, or an excessively wet or dry environment. ‘They are also able to withstand very strong sunlight and wide variations in surface tem- perature. The climax is the final or mature stage with biomass at its maximum under existing environmental conditions. There is a dynamic ‘equilibrium of nutrient inputs and outputs, Productivity, biomass, population numbers and species composition. The term ecological climax can never have absolute meaning because many factors can influence the attainment of a climax community. No environment is ever completely stable as far as physical characteristics are concerned and all ecosystems are continually adapting to changing conditions. Tt is also impossible to determine Whether an ecosystem has already reached its Chapter 3: The Dynamics of Ecosystem Functioning 51maximum stage of development under a certain sset of conditions. For example, from a climato~ logical point of view, ecosystem development may stil be possible, but is prevented by frequent fires. Active involvement by humans, and specifically commercial activities, may interfere with the attainment of the climax stage. This influence can be direct as in the case of veld fires. Under such Influences, natural succession can be controlled to create a disclimax, which would never develop under normal conditions. Regular burning can favour fire resistant species and when the limiting influence of fire Is removed, @ “normal” climax system can, in all probability, will again develop. On the other hand, practices by farmers, such as overgrazing, can lead to dete- rioration of the environment with the instability of the ecosystem as a result. Under such conditions exotic animals and plants such as Australian ‘Acacias and prickly pear may invade the area to form a disclimax due to indirect actions (see also par. 4.4.2.1). ‘The concept of succession seems simple enough 3. 10 Stability at first glance. However, in the analysis of scientific literature on the topic, it becomes clear that differences of opinion exist. One can for example query the concept of homogenous vegetation type in any so-called climax situation. Within one landscape it is perfectly normal to have grass, patches of forest, riparian vegetation and marshes - each one portraying its own climax vegetation. Some scientists discard the monoclimax idea (based on climatically con- trolled conditions) for a polyclimax idea where soil, topography, micro-climate and other envi- ronmental factors are all included. Therefore, within an area of climax vegetation, several different types of "climax vegetation" may exist. As interference by human activities in the environment increases, the number of natural ecosystems capable of reaching or maintaining an ecological climax will decrease, The chances of having the opportunity to study an ecological climax and to determine a base line for future reference, is becoming smaller and smaller. It Is therefore imperative that as many natural eco- systems as possible are conserved for the future. tability of an ecosystem refers to the ability to maintain a condition of equilibrium in species composition, biomass and productivity. In addition an ecosystem must be able to return relatively quickly to a condition of stability after it has been subjected to external disturbances. It is an accepted ecological principle that stability in an ecosystem is directly related to the number of species and to the specific composition of the populations present. This principle is based on the fact that the larger the number of species and the resultant links in the food chain, the greater the abilty of the ecosystem to overcome Changes in the external environment which can cause fluctuations in population numbers. An ‘example is the possible fluctuation in the her- bivore population if an external influence, for ‘example humans, were to eliminate carnivores lke jackals, from a simple desert ecosystem. On the contrary, the removal of a single carnivore species in a complicated ecosystem (such as removing tigers from the forest) will probably not have equally drastic consequences because there will be numerous other carnivores to TCOME 3.5 After having studied the next section you should be able to understand how dynamic balance and stability is being retained if the ecosystem.-~ control an increase of herbivores. However, this hypothesis cannot be accepted out of hand because a simple relationship between stability and the number of species is not always observed. In some cases a negative correlation exists between stability and diversity and the stability may be decreased as a result of the complex nature of certain ecosystems. Less complicated ecosystems can maintain stability for at least a short period of time in the face of environmental disturbances. 52 eEcoplon Environmental Management3.10.1 Homeostasis ‘The ability of an ecosystem to resist change and to remain in a state of equilibrium is known as homeostasis. This is attained mainly as a result of negative feedback. The maintenance of stabilty within the ecosystem as a whole is the result of the resistance or tolerance of individuals, within the system, to environmental change. Every organism has minimum and maximum limits within which it can function. These limits include environmental conditions such as temperature, height above sea level, rainfall, nutrients, etc. The closer the organism is to these limits, the more its functioning is impeded, and It will exist In 2 state of stress until completely inhibited when one of its limits are exceeded. Fig. 3.14 is an example of a lake where the fish population was able to survive over a long period of time in spite of increasing levels of sulphur 3.10.2 Dynamic equilibrium dioxide, because the pH could be controlled by the system. However, when the homeostatic ability of the system was exceeded, the pH fell drastically and large numbers of fish died. eo {50, RELEASE HY MRLION TONS.YR (Source: wins, 190) Figure 3.14: Effect of sulphur dioxide ‘on.an aquatic system 'No ecosystem can remain absolutely stable because the system is continually affected by factors from outside. As soon as an extemal factor disturbs the stability of the ecosystem, a process of restructur- ing immediately commences. A simple diagram- ‘matic representation of such adaptations is shown in Fig. 3.15. Elements closest to the disturbance will be influenced very quickly and ina most dramatic way depending on the nature of the disturbance, Further away from the disturbance the effect will be slower and less dramatic. For example, if a dam in a Nature reserve dries up temporarily, the following would possibly be observed. In the first place, all the fish would die because they are closest to the disturbance. In the second place, all the water birds that were dependent on the dam for their food, would depart for other dams in the vicinity. Competition in these dams would therefore increase, but the effect would not be drastic or immediate. In the third place the in- fluence on predators that were only partly depen- dnt on the birds at the dam will be minimal, When it rains again the dam would fill with water and ‘over time the dam would once again be repop- ulated and all life would then return to normal as it was before. However, if the dam remains dry, a ‘new system will develop on the site of the dam. A recent explosion in the numbers of penguins on Marion Island provides an interesting example of balance In the ecosystem. One explanation for the increase is related to the hunting of whales by humans. When many whales were present they ‘consumed enormous amounts of plankton leaving only limited reserves for other animals. When the whale population was depleted by commercial whaling the food network was disrupted and more food become available for fish. An increase in the fish population allowed the penguin population to increase. The result was overpopulation at the breeding sites that were trampled. Previously the king and macaroni penguins bred on a thick layer of turf and peat. This layer, up to 4m thick in places, became completely trampled and eroded so that the penguins now had to breed on bare rock. The eventual outcome of this whole process Is still uncertain, Nature has an amazing potential to recover from catastrophes. Parts of the Kuwait desert are still polluted with oil released from oil wells sabotaged during the Gulf War. In the more heavily contami- nated parts there is stil no plant life but elsewhere plants are returning. Their roots and those of crop plants in the same area are associated with a plethora of alkane-consuming bacteria - acting as bioremediation agents. ‘The process of adaptation to changing circum- stances is known as the maintenance of dynamic ‘equilibrium. Chapter 3: The Dynamics of Ecosystem Functioning 533.10.3 The vulnerability of an ecosystem Vulnerability of an ecosystem depends largely ‘on the resistance, or rather the lack of resistance, to disruptive factors of external origin. It also depends on the resilience of the system and the nature of the disturbance as well as the tolerance of individual species to changing environmental conditions (stress). Vulnerability is therefore the inability of an ecosystem to survive in the presence of disturbing factors such as climatic 3.10.4 The nature of disturbances The nature, extent, duration and frequency of disturbances will also influence the vulnerability of the system, A destructive storm in a rain forest in the Amazon Basin will only have a limited and temporary influence as the forest has already adapted to damage by storms. A traffic route that appears to cause only slight damage will, in the long run, have a far greater impact because traffic routes are alien to the environment and can cause chain reactions such as accelerated erosion, fire, colonization by allen species, the establishment of settle- ments, disruption of migration routes etc. Certain ecosystems benefit by the apparently destructive influence of veld fires, like grass- lands, while other ecosystems such as the Cape Fynbos can be ireparably damaged by too frequent fires. change, physical and chemical environmental changes, or human influences. It depends on (a) the amplitude of stability, that is, how far the system can deviate from normal before irrepara~ ble damage is caused, and (b) the elasticity of an ecosystem, that is, the rate at which the system can recover after disturbances have taken place. Figure 3.15: Adaptation within an ecosystem under stress (dynamic equilibrium) 3.1 1 Carrying capacity ( Sr uoy 3.3 Crisis over elephants in the Kruger National Park It is reported that, should the elephant population be allowed to increase unabated, there will be more than 20 000 elephants in the Park within the next 10 years. This will be devastating for the ecology as there would be a huge overpopulation, exceeding the 54 ofzopian Environmental Managementcarrying capacity by 13 000 individuals. The authorities have identified two so-called high-impact zones in the middle of the Park near Satara and Letaba, where the carrying capacity of the veld is high. Here elephants will be allowed to increase naturally. In the two low-impact zones the population will have to be decreased by 7% per year. The carrying capacity is that level or inten- sity of use that a renewable resource can tolerate and still remain productive. The con- cept of carrying capacity can never really be satisfactorily described and there are also no specific measuring techniques that do not in some way present problems. By way of illus tration the following examples of carrying capacity in a recreation resort are given. 2 Physical carrying capacity This can be measured in terms of the area per visitor, the number of caravan sites, braai areas, ablution facilities, etc. If the number of visitors exceeds the number of facilities, or the available space, then the carrying capacity is exceeded, @ Psychological carrying capacity It is exceeded when the resort becomes so congested that the quality of the recreation experience is spoiled. This carrying capacity threshold is subjective and it will also vary according to the type of the resort - depending ‘on whether it is Nature orientated or facility orientated, It is also expressed as a measure of “value for money” - keeping the experience in line with the total cost involved. ® Economic carrying capacity A minimum and maximum capacity of a resort exists without or above which, from a financial point of view, it will no longer be economically Viable. Should motor boats be allowed on a reservoir where the water is also used for drink- ing purposes, a critical point (value) will be reached when pollution levels will increase to such an extend that the cost of purifying the water, exceeds the income generated by boating. (Beeld, 2003-09-23) @ Management capacity The concept of carrying capacity and especially recreational carrying capacity has been criticized. Carrying capacity is not an objectively derived absolute reality and should be seen as a man- agement concept, which is rather judgmental, and goal orientated. Should the manager of a resort decide to make maximum profit, he could decide to pave large areas of the resort and allow people up the point where the physical carrying capacity comes into play. As an alter- native scenario the manager might wish to retain ecological integrity of the ecosystem and then he will bring as litle changes into the system as possible. Then the carrying capacity will be regarded as being very low. We can thus speak of Design Capacity. The problem of determi- ning carrying capacity has led researchers to use terminology such as Limits of Acceptable ‘Change (LAC) and Recreational Opportunity Classes (ROC) (Henderson, 1992). @ Ecological carrying capacity This is a measure of the maintenance of the biotic component of the environment. When the number of visitors to the resort causes damage to plant and animal life, the resort will begin to lose its attraction, and the threshold of the ecological carrying capacity will be exceeded. From a grazing point of view the carrying capacity is the ability of the veld to support a certain number of herbivores. Technically it can be viewed as the grazing capacity and is expressed as the area per herbivore unit per length of the grazing period. The ecological carrying capacity can also be described as the highest yield that can be maintained over a long period of time. It is a long-term concept and short-term actions must always be viewed in terms of the long term effect, There are two Chapter 3: The Dynamics of Ecosystem Functioning 55_ 3. 12 Rhythmicity variables that are difficult to predict: 1. The variation in productivity from season to season, as influenced by climatic factors. During @ good season an area can handle a greater impact than normal, while during a dry, hot season an area can be damaged by only slight use. 2. The intensity of use can be much higher for shorter periods of time as compared to sustained use over long periods. Carrying capacity encompasses a range of interwoven characteristics between the vegeta- tion, herbivores and the ecosystem. The in- herent ability of the veld to maintain a certain number of herbivores is limited by such factors as climate, especially precipitation and tem- perature, as well as the basic vegetation or veld type, for example grassland or Bushveld. On the other hand the veld type is a product of climate, geology, soil, slope, height, aspect and other factors. ‘The most important criteria used to determine the carrying capacity of an ecosystem is un- doubtedly the vegetation. Most animal species have preferences for certain plants. However, there is some overlap between species resulting in a degree of competition for the same food resources. Herbivores can be divided into three II forms of life are dependent on some form ‘of rhythmicity in Nature, such as the changes between night and day, seasons, tides, etc. Daily rhythmicity is the most common. form. Diurnal animals are active during the day and rest or sleep during the night, while nocturnal animals are active during the night. Many plants also exhibit daily rhythmic- ity: flowers may open at night but wither the following day, or they fold their leaves during the night and appear to be in a state of sleep, Rhythmicity caused by changes in light intensity is known as photoperiodism Many animals exhibit seasonal rhythmicity especially in theit mating behaviour, migr tion patterns, or changes in the feathers of birds, etc. Deciduous trees are subject to sea- sonal rhythms as are flowering plants that only main groups. Grazers (grass eaters) such as buffalo, white rhinoceros, hippopotamus, zebra and wildebeest; as well as most domestic animals; mixed feeders (grass and leaf eaters) such as elephants, impala and dulkers that live on grass, fruit, pods and leaves of trees and shrubs; and browsers (leaf eaters) which exist primarily on the leaves of trees and shrubs. Examples include, the black rhinoceros, kudu, giraffe and bushbuck. In wildlife management a distinction is drawn between grazing and browsing capacity. A knowledge of feeding preferences with regard to what is consumed, and the amount eaten, is therefore necessary in order to determine which species, the number of individuals, as well as the ratio between them, an ecosystem can maintain before there is over- grazing and deterioration. The term carrying capacity is therefore not an objectively measurable quantity. The management objectives will determine the intensity of use. Managing a game farm for tourism will require the maximum number of animals - with artificial addition of water and forage, while farming for trophy hunting will require the reduction of weaker animals to optimize reproduction of the biggest animals. To maximize meat production will on the other hand require a completely different approach to veld management and carrying capacity (see Bothma, 2002). bloom in certain seasons as well as annuals, grass, etc. The lunar rhythms (moon) also influence certain organisms, especially because of its influence on the tides. The light of the full moon also influences smaller fresh water organisms such as daphnia. Rhythmicity is generally genetically determined in most organisms. The change between day and night and between seasons only serves to synchronize the rhythmicity of the population. This synchronization can be remarkably accu- rate. In some cases a whole population of adult insects can, after months of feeding under most diverse conditions, "hatch" from the pupae stage within minutes of each other, over their entire habitat, to mate, lay eggs, and die. Adaptation or adjustment to alien conditions usually takes a long time, Birds, which are 56 © Ecoplan Environmental Managementmoved from one hemisphere to another can and Is proof that even humans are still con- moult “out of season” for a number of years trolled by inherent rhythmicity. This rhythmicity before adapting to the new conditions. "Jet is also known as biorhythms and the mech- Lag" is a condition that air travellers may éxpe- anism that controls it is called the biologi- rience when flying in an east-west direction cal clock. The environment consists of many elements that are functionally connected to each other in a dynamic way. These forces act in a very orderly way according to specific laws, principles and processes, Understanding these laws makes it possible to adapt our way of life in such a way that we can tune in with Nature and have sustainable living conditions. 1. Explain the link between li iting factors and tolerance level. 2, Is there a difference between the Law of Trigger factors and the Holocenotic principle? 3. Does one also find biological control measures amongst people? 4. Why is it so important for natural elements to Keep on re-cycling? 5. Explain the impact that the removal of vegetation will have on the water cycle. 6. _ How does the principle of Trigger factors come into play in this example? 7. In the 1970's the birds of prey were almost extinct in Europe. The same is seen happening in developing countries. What do you think is the reason for this? Chapter 3: The Dynamics of Ecosystem Functioning 57Bibliography and Further Reading Bothma, J. du P. 2002. Game ranch management. Pretoria: Van Schaik’s, Cross, A. 1990, Impoverishment of the Earth's biodiversity threatens our own survival. Geographical Journal. LC11(6):42-7. Enger, E.D. & Bradley, FS. 1995. Environmental science: a study of interre! ‘WMC Brown. jonships. Toronto: Gildenhuys, P. & Hugo, M.L. 1990/1. Die rol van indringerspesies in 'n sensitiewe ekosisteem: 'n gevalle- studie van die huiskat (Felis catus) op Marion elland. Die Suid-Afrikaanse geograaf. 18(1/2). Goldsmith, E, & Hildyard, N. 1990. The Earth report 2: monitoring the battle for our environment. London: Mitchell Beazley. Henderson, C. 1992. An extension of the recreational carrying capacity concept. Doctoral dissertation, University of Cape Town. Kok, P.D.F. et al. (Theron, G.K. & Grobbelaar, N.). 1987, Fanerogame en ekologie, Durban: Butterworth. (Odum, E,P. 1975. Ecology. New York: Holt, Rinehardt & Winston. Smith, R.L. 1990, Ecology and field biology. London: Harper & Row. White, I.D. et al. 1984. Environmental systems. Boston: Allen & Unwin. Websites Kruger Park ecosystem consists of several zones, which attract different wildlife according to the vegetation and climate: http://flyinglions.safari.co.za/Kruger-Park-eco-system.html ‘The Benguela Current Large Marine Ecosystem (BCLME) is shared between three countries - Angola, Namibia and South Africa. This website gives you an idea of an international programme of ecosystem conservatior http: //www.undp.org.za/projects/benguela.htm! This website gives some information on groundwater dependent ecosystems: http://waterhru.uzulu.ac.za/sw-gw/ecosystem_dependency.htm This website illustrates the "free" services that intact ecosystems provide for farmers and for humankind in general: hittp://www.nbi.ac.za/consfarm/pub/cfesfact-htm ‘An example of the effects of poison on the food chain: http://www. blackeagles.co.za/HelpPoisoning.htm The impact of humans on prawns in estuarine ecosystems and the effect of disturbance: bttp://zandvleltrust.org.za/art-sand%20prawns.htm| The base of the food chain in the coastal area: http://www divedudes.co.za/coral_dom.php ‘An example of what a mineral can do to the food chain: http://www.crystalflow.co.za/chlorine.htm 58 otxopion Environmental Management