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The role of herbivores in the ecosystem and management of Miombo woodlands • Heikki Roininen, Timo Veteli and Tiina Piirainen (University of Joensuu, Finland) • Research on plant animal interaction (ecology and evolution) – Insects, voles, hares, moose, reindeer and willow ptarmigan • • • Similar methods for research might be applicable for Miombo woodlands Research in Miombo and other systems may help to understand plant/animal relationships in general Presentation: to review literature on the effect herbivores in Miombo woodlands (excluding human) Reindeer and Willow ptarmigan Reindeer and willow ptarmigan browsing on willows Interactions between mammals, plants and environment (Cumming 1982) Effects of herbivores in relation to other influences (Sankaran 2005 et al.) • What are the primary factors driving the existence of Miombo (Savanna)? – Precipitation – Fire – Herbivore Relationship between MAP (Mean Annual Precipitation) and woody cover No herbivore effect on woody cover of dry savanna (MAP < 650mm) Relationship between MAP (Mean Annual Precipitation) and woody cover Two generally accepted influences in plant/animal interaction • Clear pattern between plant species richness and grazing/browsing intensity Effects on species richness of plants • Negative effect of grazing/browsing in nutrientpoor systems • BUT positive in nutrient-rich systems Comparing the influences of domestic and natural herbivores • Sustainable densities versus overgrazing – • Overgrazing and with episodic droughts has caused widespread rangeland degradation and loss of floristic and faunal diversity (reviewed by du Toit and Cumming 1999) Livestock and game in similar densities – – – – Very few studies done No difference in species richness (woody plants) Some indication that relative abundances changed In some cases game caused more evenness of woody plant species (it means higher diversity) Methods in Richardson-Kageler (2003) study • Both sites of fences (cattle or game) – Expected to be equal in the beginning – 18 cases studied (good replication!) • Densities known (equal in most cases) • More than 10 years time period • Woody plants measured Effects of livestock and game on woody plants in sustainable systems • No differences in species richness • Some indication that relative abundances of tree species changed (but similarities of tree assemblages are not analysed) • Is the effect of domestic animals equal with game? – More diverse fauna of feeders should caused more diverse tree community – More research needed! Studies on effects of large herbivores on woody plants • Ford (1987); elephants decreased higher woody plants but did not change species richness • Smart et al. (1985): species richness did not differ inside or outside of enclosures, but evenness decreased in enclosures • Cumming (1997); Heavy elephant effect (woody canopy removed). No difference in woody plant species richness less than 3 meters height • Augustine and McNaughton (2004): Native browsers (elephant and did-dik) provide important ecosystem services by suppressing rates of shrubs encroachment on commercial rangeland Herbivory on seedlings • General herbivory impacts on seedlings “not well known” (Shaw et al. 2002), mammals are causing high mortality • Invertebrates and rodents largely unstudied • Relative role of different herbivores is important to know – Observation studies; livestock and game effects overrun others • If fenced insect herbivores cause similar effects • Rodents? Study on A. drepanolobium seedling (Shaw et al. 2002) • • • • • Invertebrates more important than rodents Invertebrates and rodents together may compensate removal of large herbivores Especially removal of cotyledon increased if large mammals excluded Indirectly study shows that invertebrate community (herbivorous insect) differ on areas with mammals and no mammals Hypothetically invertebrates can have big effect – In our experiment: increase of invertebrates and there impacts inside enclosures • Rodent populations are highly cyclic or variable in rain forests, boreal and arctic zone – Difficult to demonstrate temporally variable effects (every 4th years) Historical evidence for impacts of herbivores, even aged stands of Acacia (Prins 1993) • Seedling establishment is difficult under natural browsing condition • Anthrax epidemics of impala population in Manyara NP – 1984, 1977, 1961, 1880 (rinderpest) – After these epidemics new even aged stand were established – Predictions for our experiment? Exclosure studies on the effect of (large) herbivores • In general most studies are unreplicated or poorly replicated enclosure studies in natural parks • Species richness and diversity measured, but for some reason similarities of species composition between treatments are not analysed • Usually long lasting, 10 or more years • Nothing is known about harvesting effects! What has been the role of herbivores (large) in the evolution of Miombo tree?? • Indirect evidence support the few that herbivores has been important selective factor – Defence structures, defence chemicals etc.