THÈSE - Université de Franche-Comté

THÈSE - Université de Franche-Comté THÈSE - Université de Franche-Comté

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Key-words: bottom-up regulation, grassland management, landscape ecology, population outbreaks, T. europaea. Introduction In temperate Europe, intensive and innovative agriculture has led to the creation of new habitats, which are suitable for small mammals (Robinson & Sutherland 2002). In many cases, this has resulted in the magnification of normal population fluctuations and significant crop damage. Rodents in general and voles in particular cause major economic losses through damage to crops (Jacob 2003). In Franche-Comté (France), a dramatic outbreak of the water vole Arvicola terrestris in 1998 destroyed 60 % of the annual forage yield, which represented a loss of € 38 million over an area of 120 000 ha (Service Régional de la Protection des Végétaux, pers. com.). Habitat manipulation by land use management could improve strategies of rodent pest control. This therefore requires a sound understanding of the population dynamics of the species concerned. Population dynamics of small mammals may be characterized by seasonal and pluriannual periodic fluctuations in density with four successive phases, as follows: low density, increase, high density and decline (Krebs & Myers 1974; Taitt & Krebs 1985). Early studies described a large diversity in population dynamics and made a number of explanatory hypotheses (Lindström et al. 2001; Hansson 2002). In arctic and boreal ecosystems, cyclic population dynamics were considered to be regulated mainly by predation (Korpimaki & Krebs 1996; Gilg, Hanski & Sittler 2003) and habitat productivity (Jedrzejewski & Jedrzejewska 1996; Ekerholm et al. 2004). In the more complex temperate ecosystems (Central and Western Europe), population cycles are not caused by one single factor (Hudson & Bjornstad 2003) and should be explained by the combined action of a hierarchy of many regulating factors in space and time (Lidicker 1995; Lidicker 2000; Hansson 2002). The abundance of small mammals seems to be principally regulated by top-down forces (predation, parasitism), and bottom-up forces (ressources) (Lindström et al. 2001; Korpimaki et al. 2004). Indication for top-down regulation was obtained on a regional scale (n x 50 km) and on a sectorial scale (n x 5 km) where the landscape may be a filter for prey/predator relationships and thus may indirectly control rodent population dynamics (Delattre et al. 1996). Earlier studies have also Thèse C. Morilhat 2005 99

shown that farming practices can impact small mammal populations. For instance, Jacob et al. (2001; 2003) demonstrated a gradient of short-term farming practice disturbances (mulching < mowing < harvesting < grazing < harrowing < ploughing) on the common vole Microtus arvalis populations (Jacob & Halle 2001; Jacob 2003; Jacob & Hempel 2003). Edwards, Crawley & Heard (1999) showed that farming practices such as grazing, soil acidification and plant removal could reduce molehill production in grassland parcels by reducing the availability of mole Talpa europaea principal food source (earthworms). In eastern France and in western Switzerland, cyclic population outbreaks of the fossorial form of the water vole Arvicola terrestris scherman (Shaw 1801) have been observed on a large scale since the 1970s in farmland, mainly in mid-altitude mountains (Jura, Massif Central, Alps) (Saucy 1994; Giraudoux et al. 1997; Fichet-Calvet et al. 2000). In these locations, population outbreaks are travelling waves, starting from clusters of communes (French administrative units) called “epicentres”, where high density occurs first, and spreading to “diffusion” communes on a 6-years cyclic pattern. High population density on a large extent causes severe crop damage and important economic losses (Meylan 1981). They have also been linked to a higher prevalence of human alveolar echinococcosis, a lethal parasitical disease maintained via a fox - small mammal cycle (Raoul et al. 2001). Chemicals have been used to control water vole population outbreaks but they may result in secondary poisoning of non-target wild animals (Brakes & Smith 2005). Evidence has been provided that the risk of population outbreak is related to landscape features, e.g. the ratio of Permanent Grassland (optimal habitat for the water vole) to Agricultural Land (PG/AL), on the regional scale. This has been termed: the landscape composition effect (Giraudoux et al. 1997; Fichet-Calvet et al. 2000). The authors also showed that population destabilization occurs when a particularly high PG/AL ratio is found (> 85%). On a sectorial scale, Duhamel et al. (2000) showed that epicentres are generally comprised of large grassland openfields (51 %) and few forest (22 %), as compared to diffusion communes (respectively 32 and 46 %). This has been termed the landscape structure effect. On a parcel scale (n x 10 m) two earlier studies, carried out in the farmland of the Swiss Jura mountains, suggested a bottom-up regulation of water vole populations. Capture-markrecapture and radiotelemetry experiments indicated that the first vole colonies were distributed in grassland parcels with abundant plant cover (Saucy 1988). In a long-term study, on the Thèse C. Morilhat 2005 100

shown that farming practices can impact small mammal populations. For instance, Jacob et al.<br />

(2001; 2003) <strong>de</strong>monstrated a gradient of short-term farming practice disturbances (mulching <<br />

mowing < harvesting < grazing < harrowing < ploughing) on the common vole Microtus<br />

arvalis populations (Jacob & Halle 2001; Jacob 2003; Jacob & Hempel 2003). Edwards,<br />

Crawley & Heard (1999) showed that farming practices such as grazing, soil acidification and<br />

plant removal could reduce molehill production in grassland parcels by reducing the<br />

availability of mole Talpa europaea principal food source (earthworms).<br />

In eastern France and in western Switzerland, cyclic population outbreaks of the fossorial form<br />

of the water vole Arvicola terrestris scherman (Shaw 1801) have been observed on a large<br />

scale since the 1970s in farmland, mainly in mid-altitu<strong>de</strong> mountains (Jura, Massif Central,<br />

Alps) (Saucy 1994; Giraudoux et al. 1997; Fichet-Calvet et al. 2000). In these locations,<br />

population outbreaks are travelling waves, starting from clusters of communes (French<br />

administrative units) called “epicentres”, where high <strong>de</strong>nsity occurs first, and spreading to<br />

“diffusion” communes on a 6-years cyclic pattern. High population <strong>de</strong>nsity on a large extent<br />

causes severe crop damage and important economic losses (Meylan 1981). They have also<br />

been linked to a higher prevalence of human alveolar echinococcosis, a lethal parasitical<br />

disease maintained via a fox - small mammal cycle (Raoul et al. 2001). Chemicals have been<br />

used to control water vole population outbreaks but they may result in secondary poisoning of<br />

non-target wild animals (Brakes & Smith 2005). Evi<strong>de</strong>nce has been provi<strong>de</strong>d that the risk of<br />

population outbreak is related to landscape features, e.g. the ratio of Permanent Grassland<br />

(optimal habitat for the water vole) to Agricultural Land (PG/AL), on the regional scale. This<br />

has been termed: the landscape composition effect (Giraudoux et al. 1997; Fichet-Calvet et al.<br />

2000). The authors also showed that population <strong>de</strong>stabilization occurs when a particularly high<br />

PG/AL ratio is found (> 85%). On a sectorial scale, Duhamel et al. (2000) showed that<br />

epicentres are generally comprised of large grassland openfields (51 %) and few forest (22 %),<br />

as compared to diffusion communes (respectively 32 and 46 %). This has been termed the<br />

landscape structure effect.<br />

On a parcel scale (n x 10 m) two earlier studies, carried out in the farmland of the Swiss Jura<br />

mountains, suggested a bottom-up regulation of water vole populations. Capture-markrecapture<br />

and radiotelemetry experiments indicated that the first vole colonies were distributed<br />

in grassland parcels with abundant plant cover (Saucy 1988). In a long-term study, on the<br />

Thèse C. Morilhat 2005 100

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