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Return of the Wolves Implications of the Role of Top Predators in Terrestrial Ecosystems for the Return of Eurasian Wolf Canis Lupus Lupus to Western Europe

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Return of the Wolves Implications of the Role of Top Predators in Terrestrial Ecosystems for the Return of Eurasian Wolf Canis Lupus Lupus to Western Europe Anne M.G. Kwak Return of the wolves Implications of the role of top predators in terrestrial ecosystems for the return of Eurasian wolf Canis lupus lupus to Western Europe Author Anne M.G. Kwak Radboud University Nijmegen E-mail: [email protected] Education Master Biology, track Communities & Ecosystems Radboud University Nijmegen Faculty of Science Heyendaalseweg 135 6525 AJ Nijmegen Phone: 024-365 26 61 Fax: 024-355 34 50 Website: www.ru.nl/fnwi Supervisor Prof. dr. H. (Hans) de Kroon Institute for Water and Wetland Research Radboud University Nijmegen Date October 2013 SUMMARY In many terrestrial ecosystems in Western Europe today, top predators are absent (Terborgh et al., 1999). Since top predators are considered to be important for the structure of ecosystems and their biodiversity, the absence of such species might have large impacts on these systems. In this thesis I want to investigate how the absence of top predators has affected terrestrial ecosystem trophic structure and biodiversity. The focus on terrestrial ecosystems comes forth from the fact that the Eurasian wolf Canis lupus lupus is currently expanding its range throughout Western Europe (Chapron et al., 2003; Randi, 2011). Here, I investigate how species like the Eurasian wolf might affect our ecosystems when they spread further, and what the implications of these effects are for the management of our ecosystems and of the Eurasian wolf. Carnivores can be assigned the role of keystone species, where they create ecological boundaries that protect lesser competitors from exclusion (Miller et al., 2001). In this role of keystone species, carnivores are also capable of increasing plant biomass through their limiting effect on herbivore numbers, causing both plants and carnivores to flourish (Miller et al., 2001). A general pattern in terrestrial ecosystems seems to be that when top predators are absent, herbivores increase in population size, resulting in a high browsing pressure (Elmhagen and Rushton, 2007; Miller et al., 2001; Ripple and Beschta, 2004; Ripple and Beschta, 2006). This eventually leads to a lower biodiversity than in systems where predators are still present (Sergio et al., 2005). Ripple and Beschta (2006) also found that removal of top predators does not only affect the direct food chain in which the predator is present, but also other parts of the ecosystem. So what effects can Eurasian wolves Canis lupus lupus have on ecosystems? First of all, since they are top predators, Eurasian wolves can contribute to the regulation of predator-prey relationships. Next to this they also affect the density of smaller predators (Randi, 2011). The expansion of the Eurasian wolf to western Europe may provide us with a way to passively manage and restore our ecosystems to a more natural and wild state. In each of the example areas from Europe, a wolf population will result in different effects on the present ecosystems, but it is still expected to lead to an increased biodiversity. It would also reduce the need for human interference in these systems, leading to more natural states. However, top predators like the Eurasian wolf do not only bring with them changes to ecosystems, but also evoke fear in humans either for their own safety or for that of their domestic and game animals. Management and conservation of large carnivores may thus only be successful in the long term if people can accept free ranging predators in their area. To achieve this, we will have to regulate the wolf population to address public concern, but at the same time we need to maximize population viability of these predators (Chapron et al., 2003). To conclude, the Eurasian wolf returning to Western Europe may have very positive effects from an ecological perspective: increased biodiversity. However, we do need to consider public fear. We will have to come up with management strategies now, to build up support from the public in order to welcome back the wolf to our ecosystems. CONTENTS Summary 1 1 Introduction 5 2 Top-down versus bottom-up regulation of ecosystems 6 2.1 Food webs and keystone species 6 2.2 Bottom-up versus top-down ecosystem regulation 7 3 Absence of top predators 10 3.1 Importance of top predators in terrestrial ecosystems 10 3.2 Effect on the structure of ecosystems and food webs 11 3.3 Management of ecosystems lacking top predators 13 4 Current conservation issues 15 4.1 Implications for management of Eurasian wolf Canis lupus lupus 18 5 Discussion 20 6 Literature 21 MSc thesis Return of the wolves Anne M.G. Kwak 1 INTRODUCTION There has been an ongoing discussion on top-down versus bottom-up ecosystem regulation since the 1960s (Elmhagen and Rushton, 2007; Power, 1992; Terborgh et al., 2001), when the concept of top- down regulation was first introduced by Hairston et al. (1960). The question is whether ecosystems are regulated from the top down by predators or from the bottom up by food/nutrients limitations. Most evidence for top-down regulation in ecosystems comes from aquatic ecosystems (Elmhagen and Rushton, 2007). However, mammalian predators are considered to have major influences on the trophic structure and biodiversity of terrestrial ecosystems (Hebblewhite et al., 2005; Sinclair et al., 2003; Terborgh et al., 2001). In many terrestrial ecosystems in Western Europe today though, top predators are absent (Terborgh et al., 1999). Such ecosystems are often managed by humans to keep them in a certain state or to at least preserve a certain level of biodiversity. Since top predators are considered to be important for the structure of ecosystems and their biodiversity, the absence of such species might have had large impacts on these systems. In this thesis I want to investigate how the absence of top predators has affected terrestrial ecosystem trophic structure and biodiversity. The focus on terrestrial ecosystems comes forth from the fact that Eurasian wolf Canis lupus lupus is currently expanding its range throughout Western Europe (Chapron et al., 2003; Randi, 2011). For example, for the first time in 150 years a wolf has been found in the Netherlands (NU.nl/ANP, 2013). Here, I investigate how species like the Eurasian wolf might affect our ecosystems when they increase in numbers, and what the implications of these effects are for the management of our ecosystems and of the Eurasian wolf. To answer this question, I will first introduce the concepts of top-down and bottom-up ecosystem regulation and investigate whether or not a consensus has been reached within the discussion on which mechanism is most important. Next, I will investigate the effect of absence of top predators on terrestrial ecosystems to see how trophic structure and biodiversity have altered. Finally, I discuss the consequences of the expansion of Eurasian wolf to Western Europe on our ecosystems and I will link this to the implications this expansion has for management of this species and our systems. Radboud University Nijmegen | 5 MSc thesis Return of the wolves Anne M.G. Kwak 2 TOP-DOWN VERSUS BOTTOM-UP REGULATION OF ECOSYSTEMS 2.1 FOOD WEBS AND KEYSTONE SPECIES Ecosystems can be described as food webs: several food chains linked together into a complex web of interactions between organisms (Begon et al., 2003; Jordán, 2009; Jørgensen, 2009). Using a food web approach allows us to investigate the importance of different species from an ecological perspective. While such food webs can be very large and complex, a ‘small-world’ structure seems to be common in nature: most species only interact with a small number of other species, while the connectivity of food webs as a whole is maintained by a few species that interact with a large number of other species (Jørgensen, 2009). Extinction of such major interactors may break coevolved relationships among co- adjusted sets of species and may thus significantly harm ecosystem integrity (Jordán, 2009). This concept of a ‘small-world’ with major interactors forms the basis for the idea of keystone species: small-world networks are robust to random loss of nodes (species), but they are vulnerable to attacks targeting highly connected nodes (keystone species) (Jørgensen, 2009). When looking at the interactions between species present in food webs, one might have an initial idea of its workings. One would expect removal of predators to lead to an increase in prey density, or removal of a competitor species to lead to an increase in abundance in the species it competes with. However, this is not always the case. Occasionally, removal of a predator leads to a decrease in prey species or removing a competitor causes a decrease in the abundance of its competitors. These effects occur when direct effects in a food web are less important than indirect effects (Begon et al., 2003). These indirect, ‘unexpected’ effects are a common occurrence in nature. They refer to the propagation of perturbations through one or more trophic levels, so that consequences are felt in organisms that may seem far removed from the subjects of perturbation (Begon et al., 2003; Terborgh, 1988). Species that show such effects in food webs upon their removal are usually keystone species (Begon et al., 2003). The term ‘keystone species’ was first introduced in 1969 by R.T. Paine as a species which presence is crucial in maintaining the organization and diversity of their ecological communities which makes it implicit that they are exceptional in their importance (Mills et al., 1993). They show disproportionally large effects relative to their abundance and are by definition single objects on which the forces keeping a community intact are concentrated (Payton et al., 2002). Removal of keystone species leads to extinction or large changes in abundance of several species (both through direct and indirect pathways), resulting in a community with a very different species composition and, to us, different physical appearance (Begon et al., 2003).
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