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The Modification Process of Sika Deer Impact on Forest Vegetation by Artificial Grasslands

財部, 薫乃子 筑波大学

2020.07.22

概要

Ungulates (e.g., deer) can profoundly alter the structure and composition of forest communities via both direct and indirect mechanisms (Rooney and Waller 2003). Selective ungulate herbivory leads to the dominance of unpalatable species and/or tolerance-trait species in communities, which have chemical defense mechanisms or regrowth capacity (Augustine and McNaughton 1998). For instance, through experimental manipulation of white-tailed deer (Odocoileus virginianus) density, Nuttle et al. (2014) demonstrated that higher deer densities caused increased fern domination in the understory, decreased seedling number and forb cover, and diminished angiosperm richness. On the other hand, it is also known that ungulate herbivory on individual plants mitigates interspecific competition among plants, leading to high species diversity (Nishizawa et al. 2016). Moreover, Vild et al. (2017) showed that ungulate herbivory significantly increased α-and γ-diversity, and which caused significant vegetation homogenization inside the game preserve, and which caused by massive enrichment with ruderal species. In the Arctic, higher reindeer (Rangifer tarandus) densities have been linked to decreased plant species richness in less productive sites and to increased species richness in more productive sites (Sundqvist et al. 2018).

Many studies have suggested that ungulate herbivory affects forest regeneration and that the impact of the deer is driven by various factors. Cretaz and Kelty (2002) revealed that dense fern (Dennstaedtia punctilobula) understories showed differential interference among species with seedling development after deer browsing was reduced. Uno et al. (2019) showed that deer browsing had a direct negative effect on seedling survival as well as the indirect positive effect of reducing bamboo (Sasa senanensis) coverage in a cool-temperate mixed forest on Hokkaido. Murata et al. (2009) reported much greater negative effects of Sasa borealis than of sika deer browsing on seedling emergence and survival in cool-temperate mixed forests in Kyushu. Akashi et al. (2011) demonstrated that percentage of browsed seedlings was positively correlated with deer abundance, and that it was affected by deer seedling species preference in Abies sachalinensis plantations in Hokkaido. In addition, Gill and Beardall (2001) showed that deer greatly contribute to the seed dispersion of many plant species, and that plants with small, hard seeds are most likely to survive digestion. Hence, the effects of ungulate herbivory occur at different hierarchical scales, ranging from individual plants to whole forest ecosystems.

Deer have expanded their range and have increased dramatically in abundance worldwide in recent decades. Their impact on natural ecosystems has been accordingly dramatic (Côté et al. 2004). In Japan, the population size and range of sika deer (Cervus nippon Temminck) have been rapidly increasing in many regions since the 1970s, and the impact of their herbivory on forest vegetation has increased remarkably (Takatsuki 2009). Hence, sika deer population control has been conducted to decrease the damage on agricultural production and forest ecosystems. Sika deer are medium to large ungulates native to Southeast Asia, eastern China, and Japan (Kalb et al. 2018), and they have become problematic in many regions in which they were introduced as novelties or games species as they hybridize with or are aggressive toward native deer species (Germany and Austria; Pitra et al. 2005, New Zealand; Husheer et al. 2006, USA; Kalb et al. 2018). Husheer et al. (2006) suggested that because of their dietary advantage, introduced sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites in which impacts on tree regeneration are likely to be greatest.

Deer impact on forest vegetation is driven by various factors, such as deer density (Côté et al. 2004), individual species characteristics (Horsley et al. 2003), light conditions (Suzuki and Ito 2014), snow depth (Kiffner et al. 2008), and so on. Recently, landscape composition has attracted attention as an important factor that strongly affects the impact of deer herbivory on forest vegetation. However, there are limited and inconsistent studies that have evaluated the degree to which browser impact on vegetation is scale-dependent on variation in ungulate density and habitat composition (Royo et al. 2017). Some studies have shown that herbivory intensity on the forest understory becomes lower in fields with rich food, such as pastures near to forests, than in fields without rich food (Takada et al. 2002; Honda et al. 2008; Royo et al. 2017). On the other hand, some studies have demonstrated that food-rich habitats increase the deer impact on surrounding forest (Reimoser and Gossow 1996; Iijima and Nagaike 2017). Therefore, it is important to generalize the process of how artificial landscapes affect deer impact on forest vegetation in order to understand forest dynamics.

Among various food-rich habitats, I focused on artificial grasslands, such as pastures, because they can provide a large quantity of high-quality and digestible grasses for deer (Takatsuki 2001). Previous studies have reported that deer often forage grasses in artificial grasslands (Trdan and Vidrih 2008; Kamei et al. 2010; Iijima 2018). Moreover, it has been suggested that artificial grasslands increase the population growth rate (Iijima et al. 2013) and the carrying capacity of sika deer (Iijima and Ueno 2016). In Japan, the area of artificial grasslands increased from 1965 (139,800 ha) to 1994 (661,400 ha). Since then, it has been declining and reached its lowest point (603,400 ha) in 2016 (MIAC 2016). Although there is some debate due to the complexity of distinguishing artificial grasslands from other landscape components (Matsuura 2016), previous studies have suggested that artificial grasslands currently occupy 1%– 5% of Japan’s total land area (Ogura 2006, Matsuura et al. 2012). At first glance, this figure seems very low, but, considering the strong influence of artificial grasslands on sika deer, its effects should not be ignored.

Hence, to clarify the processes that modify how sika deer impact forest vegetation as a result of artificial grasslands, I conducted the following set of studies:
1. I aimed to clarify how artificial grasslands around forests affect the deer density in the forests and their impact on forest vegetation while considering the plant species at spatially finer scales (Chapter 2). Specifically, I examined whether 1) the surrounding abundant alternative food (in this study, artificial grasslands) in forests increase deer density and 2) the high deer density causes the more serious impact on forest vegetation. This chapter is in press.
2. I aimed to clarify the effect of presence/absence of artificial grasslands on sika deer (Cervus nippon) impact on forest vegetation (Chapter 3). I compared the occurrence of debarking and the coverage of understory vegetation in two adjacent regions under similar deer densities with and without artificial grasslands in Yamanashi Prefecture, Japan. This chapter was published in Takarabe and Iijima (2019).
3. I aimed to clarify the effect of presence/absence of artificial grasslands on the seasonal food habits of sika deer (Chapter 4). I conducted a pellets analysis to evaluate the seasonal food habits of sika deer in two adjacent regions under similar deer densities with and without artificial grasslands in Yamanashi Prefecture, Japan.
4. The final chapter contains a general discussion regarding the effects of artificial grasslands on deer impact on forest vegetation in a cool-temperate forest (Chapter 5).

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