Biodiversity related to running waters PD Dr. Carsten Hobohm from University of Lueneburg, Germany Lecture 4/2002 in Innsbruck, Austria Summary Many organisms are adapted to running waters. Some of them live within the water for their whole lifecycle. Because of disturbances or unfavorable seasons in running waters dispersal over long distances is advantageous for plant and animal species living in rivers. Worldwide only few species are endemics of only a single river. Some examples of plant and animal species diversity, of endemic and non-endemic species and their relationship to different habitats related to running waters are discussed. Biodiversity Life and its extraordinary diversity is the unique wealth which distinguishes the plant earth from all other planets in the universe. Mankind is a critical element of this wondrous spectrum, and thus biodiversity in all its aspects represents the very foundation of human existence. We have only become aware of the real dimensions of this wealth during the last thirty years, and today we know some 1.7 million different species probably representing far less than 10 or 20 % of the actual diversity. At the same time it is increasingly evident that - due to the rapid growth of the world`s population which is about six billion people and is driving the extreme exploitation of natural resources - this diversity is undergoing a dramatic change. Fundamental genetic resources, which evolved over more than three billion years, probably are being lost at a dramatic pace. Many quantitative approaches to biogeography have been published (see e.g. Barthlott & Kier 2001, Barthlott & al. 1999, Coleman 1981, 1982, Connor & McCoy 1979, Hobohm 2000a, Iwasa & al. 1994, Malyshev & al. 1994, Malyshev 2000, MacArthur & al. 1966, MacArthur & Wilson 1967, Myers & Giller 1988, Simberloff 1974, Whittaker 1972, Wilson & Shmida 1984). The numbers of species and endemics of many groups of organisms are well known in many parts of the world, for example the numbers of vascular plant and mammal species in individual countries or national parks. According to Mittermeier & al. (1999) and Hobohm (2002) many biodiversity hotspots are concentrated in the tropics. E.g. mammals, birds, reptiles, amphibians, insects and vascular plants are show high concentrations in the tropics. However, until now neither the function of water, especially of running waters, for biodiversity nor the relationship between biodiversity and running waters has been analyzed in detail. It is impossible to explain or discuss the systematics, ecology or foodweb of rivers in a single lecture. Therefore the following topics will bring only examples which can show the complexity of biodiversity related to running waters. Species richness and ecosystem processes - theoretical relationships Diversity would not be interesting if the level of diversity were the same everywhere. Fortunately, no matter how diversity is defined, or what types of organisms are being considered, there is a phenomenal variation in diversity across the entire range of living systems. No single process or theory can explain a phenomenon as complex as biological diversity. The intellectual challenge and scientific value of the study of diversity lies in the conceptual synthesis required to understand a complex phenomenon that is influenced by many different interacting factors and processes. Theory and data have, until very recently, focussed - at least very often only - on the relationship between species richness and ecosystem processes. Taking this state of affairs as our starting point, within any one site or location, we can imagine ecosystem processes responding in one of three ways to reductions in species richness. 1. The redundant species hypothesis suggests that there is a minimal diversity necessary for ecosystem functioning, but beyond this minimum, most species are redundant in their roles; above this minimum, adding or deleting species has no detectable effect on the process or processes in question. 2. In marked contrast to (1), the rivet hypothesis postulates that all species are important (by analogy with the rivets holding an aeroplane together, so that ecosystem processes are progressively more impaired as species are lost from the system. If some rivets are missing the aeroplane is able to fly. 3. The idiosyncratic hypothesis again postulates that species are important in ecosystem processes, but particular species identities matter more than species richness per se. In consequence, ecosystem processes change erratically and unpredictably as species are lost from the system. Providing the species pool is reasonably large (thereby avoiding strong stochastic effects), the most likely theoretical relationship between loss of species and ecosystem processes is pattern, consistent with the rivet hypothesis. The fundamental underspinning mechanism is that species differ in their ecologies, that is, no two species have identical niches. But there are some important, and still poorly understood, subleties underlying this simple statement. 4. The sampling hypothesis. Species differ intrinsically in their potential maximum size, growth rate, and so on. Hence, mixtures of many species are more likely to contain high-yielding species than monocultures or low-diversity mixtures. In consequence, random samples of species from a pool will, on average, show higher biomass, and higher productivity, as species richness increases. 5. The niche complementary hypothesis. Here the niche differences are explicit. Niche-differences between species ensure that as species richness increases in ecological assemblages, so does the range of functional spce occupied by the assemblage. More diverse communities are more spacefilling above ground, have a wider range of requirements for ressources, and so on. 6. The posive species interactions hypothesis. Species do not compete with each other in communities. It is becoming increasingly clear that some plant species for example benefit from the presence of other species in the community. These hypotheses are good for understanding ecological conditions and species compositions in an ecosystems. But ecosystems cannot exist by themselves. For a good understanding we do need the knowledge of some dimensions more. In general, three groups of mechanisms are important: * mechanisms of evolution * processes which provide dispersal * ecological processes time species diversity of a region processes of evolution (evolutionary times) ecological mechanisms (ecological times) dispersal (intermediate times) area fig. dimensions of species diversity Biodiversity and running waters in space I: Biodiversity in running waters There is a foodweb in every zone of a river which is connected with other foodweb structures in the semi-terrestrial and terrestrial surroundings and other zones of the river. This complex is part of different other lectures of the IP. As a result, it will not been discussed here. Biodiversity and running waters in space II: Biodiversity close to the river (riverside) Biodiversity related to running waters is not only the biota of running waters. There are a lot of biotopes directly connected with river systems. Different habitats are typical parts of riversides in Europe. They can be ordered in different ways: * from wet (small lakes) to dry (dunes, rocks) * according to biomasses, pure substrates (riverbanks) to forests (e.g. with Salix spp.) * according to human activities from untouched to artificial and so on. Biodiversity and running waters in space III: Biodiversity indirectly dependent on rivers Many plant and animal species profit from the existance of rivers, but they are not connected with or directly dependent on rivers. For example many storks are breeding near large rivers. They are feeding in many biotopes, e.g. wet places, but also in cultivated areas outside river-valleys. Many other birds use rivers for orientation as traffic lines. Biodiversity and running waters in time The evolution, also the biology of dispersal, population dynamics in different groups of organisms and in different limnic and terrestrial habitats, depends on the quality and quantity of running waters in space and time. Biodiversity and running waters in time I: ecological times Biodiversity and running waters in time II: dispersal Because of disturbances or unfavorable seasons in running waters dispersal over long distances is advantageous for plant and animal species living in rivers. The result are highly adapted organisms living in or close to running waters. Comparing the species numbers of limnic and terrestrial organisms, rivers are poor. This is an effect of area sizes, age and unfavorable conditions. Lands are larger and older than rivers - thinking in geological times. As a result unfavorable conditions, e.g. drought or negative temperatures can extinct whole populations in rivers whereas it is more easy to survive in large terrestrial ecosystems. On the other hand, e.g. terrestrial vascular plant species in Central Europe seem to be concentrated where large rivers come together. This fact is not easy to understand, because habitat diversity very often is contradictory in these landscapes. Biodiversity and running waters in time III: evolutionary times Compared with terrestrial ecosystems most rivers are very young. Within running waters worldwide not many species are endemics of only a single river. But e.g. some fishes or river-dolphins (Inia geoffrensis in Amazonas, Lipotes vexillifer in the yang Tse, Plantanista minor in the rivers Ganges, Brahmaputra and Meghna, Platanista gangetica in river Ganges) are endemics in some large and old river-systems. Some plant species, e.g. Oenanthe conioides in the Elbe river-valley, are also endemics related to running waters. Running waters can devide terrestrial lands in different parts. For example many organisms, e.g. many small mammals, reptiles or amphibians, are not able to cross large rivers. Separation is one possibility for initiating genetic isolation. Genetic isolation on the other hand can result in adaptive radiation. So evolution can be stimulated by separation. References Arrhenius O (1921) Species and area. - J. 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