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Seasonal Impact of Deer Browsing on the Demography of Fagus Japonica Seedlings in a Cool-Temperate Forest

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Seasonal Impact of Deer Browsing on the Demography of Fagus Japonica Seedlings in a Cool-Temperate Forest 日緑工誌，J. Jpn. Soc. Reveget. Tech., 39（2）, 232―240,（2013） 論文 ORIGINAL ARTICLE SPECIAL ISSUE “Protection and restoration of vegetation damaged by deer grazing” Seasonal impact of deer browsing on the demography of Fagus japonica seedlings in a cool-temperate forest ISHIZUKA, Wataru＊1）,KAJI,Mikio2） and GOTO, Susumu2） 1）Graduate School of Arts and Sciences, the University of Tokyo 東京大学大学院総合文化研究科 2）Graduate School of Agricultural and Life Sciences, the University of Tokyo 東京大学大学院農学生命科学研究科 Abstract: Severe browsing pressure from deer may have great effects on the regeneration dynamics of woody seedlings by affecting seedling mortality and seedling demography. This study quantified the seasonal impact of deer browsing on the relationship between seedling survival and the timing of seedling germination. We es- tablished a fenced deer exclosure in a cool-temperate forest in Chichibu that experienced browsing pressure from deer. Newly germinated seedlings of the dominant beech, Fagus japonica, were marked and monitored frequently in the fenced plot and outside the deer fence (open plot). To compare the demography of current- year seedlings, the effects of the timing of germination, light conditions, and total seedling density in each moni- toring quadrat were analyzed considering their differences between the two types of plots. The survival rate at the end of the growth period was significantly lower in the open plot (2.0％)thaninthefencedplot(9.4％), mainly because of deer browsing (73.8％ of the total mortality in the open plot). Intensive browsing was ob- served only during the early growth season, while damping off was the major cause of mortality in the fenced plot (79.0％ of total) during the middle season. Statistical analysis detected opposite effects in the open and fenced plots; early germination was disadvantageous and advantageous for survival, respectively. Seedling de- mography relating to the advantage of the timing of germination may change depending on the impact of deer browsing, which has clear seasonality. Key word: early emergence advantage, deer browsing, Fagus japonica, seedling demography 石塚 航・梶 幹男・後藤 晋：冷温帯林におけるイヌブナ実生動態への季節的なシカ摂食害の影響 摘要：シカの高い摂食圧は樹木の実生生残数のみならず，生残条件をも変え，更新動態に大きな影響を与えると みられる。本研究では，実生発生タイミングと当年生残との関係における，シカの季節的な摂食害の影響を評価 するため，シカ害のみられる秩父地方の冷温帯林にてシカ防護柵を設置し，柵内外の調査枠内に発生した優占種 イヌブナの当年生実生を調査対象とした。個体識別して実生の消長を追跡し，当年生残に対する発生タイミング の効果を，生育地の光条件や実生密度の影響とともに分析し，柵内外で比較した。柵外区の実生当年生残率（2.0 ％）は柵内区の生残率（9.4％）よりも有意に低く，死亡実生の 73.8％ がシカ摂食害によるものだった。死亡要 因は顕著な季節性を示し，シカ害は実生発生初期に，柵内区の主たる死亡要因である菌害（全実生の 79.0％） は生育中期に集中していた。統計解析によって，柵内外で実生の発生タイミングが当年の生残に影響しているこ とがわかったが，柵外区では早い発生が不利に，柵内区では早い発生が有利になっており，対照的な影響が検出 された。早い発生の有利性はシカの存在下では失われ，季節性のあるシカの摂食によって実生生残の動態が変わ ることもあると示唆された。 キーワード：早い発生の有利性，シカ食害，イヌブナ，Fagus japonica，実生生残過程 is an important and well-known issue3，26，30）. Many studies 1. Introduction have shown that deer significantly affect the survival of rela- The impact of deer browsing on various forest ecosystems tively large current-year seedlings by direct brows- ＊Corresponding author：3―8―1 Komaba, Meguro-ku, Tokyo, 153―8902 Japan E-mail：[email protected] Seasonal impact of deer browsing on the demography of Fagus japonica seedlings in a cool-temperate forest 233 ing4，16，14，20，24，30，31）. In cool-temperate forests, low seedling re- one of the dominant species in climax forest, together with cruitment because of increased deer populations may affect Fagus crenata Blume, which is another Japanese beech spe- forest regeneration negatively14，19）. Therefore, deer browsing cies, and a coniferous tree, Tsuga sieboldii Carrière. These is regarded as a severe limiting factor for seedling survival two beech species are well-known of exhibiting a habitat of in these forests. However, seedling germination and subse- mast seedling at intervals of several years. Mature seeds are quent establishment are the most vulnerable life stages for mainly dispersed by gravity and germinate during the next temperate forest trees6，12）. Therefore, seedling survival for spring. After germination, the seedling establishment proc- such trees is affected strongly by deer browsing, damping ess of beech is strongly affected by multiple limiting factors, off, small mammals, and other predators such as in- especially during the initial growth period. Seedling mortal- sects1，22，27）. ity seems to be high under a closed canopy or dwarf bam- In addition to those mortality factors, the conditions of boo layer because of their light demand and susceptibility to each seedling may also affect its survival, e.g., the germina- pathogenic fungi. Predation by small mammals, other verte- tion phase, light conditions, and vegetation density. Several brates, and insect larvae also increase the mortality of beech studies have demonstrated an advantage of early germina- seedlings. Therefore, the seedling recruitment of F. japonica tion on seedling survival over late germination1，2，10，27，32）.Inde- is highly limited in natural stands8）. ciduous forests, the forest canopy becomes closed during 2.2 Study area and the two experimental plots leaf flushing; therefore, there is a seasonal limitation on the Our study was conducted on the southwestern steep slope available resources in the understory (i.e., ephemeral light). of Mt. Hakutai (1,794 m a.s.l.) in the University of Tokyo Early germinating seedlings, therefore, can utilize these lim- Chichibu Forest (35°55′N, 138°50′E, 1,210―1,260 m a.s.l., with ited resources efficiently and grow sufficiently during the a slope inclination of approximately 33°), which is located in early growing season28）. An advantage of early germination central Japan. The mean temperature and annual precipita- may be detected if the timing of seedling germination is re- tion between 1993 and 2001 at the nearest meteorological lated to the resistance to mortality caused by pathogens station (200 m apart from the study area on 1,160 m a.s.l.) and／or the preferences of predators such as small ro- were 8.9°C and 1,416 mm, respectively. This region belongs dents1，13，27）. Madsen reported17） that emerged beech seed- to the cool-temperate zone and is categorized as having a Pa- lings (Fagus sylvatica L.) were browsed immediately by cific Ocean-type climate because there is more precipitation deer, which suggests that the advantage of early germina- in the summer than in the winter. The study area was cov- tion may not be maintained if deer browse severely. How- ered by an old growth forest with a total basal area of 48.7 ever, the relationship between the effect of deer browsing m2／ha, and no management activities were done to facilitate and the timing of germination on seedling survival remains monitoring the ecosystem during long-term research. The unclear. relative abundances of three dominant species, F. japonica, Observational study was, then, conducted to trace the de- F. crenata,andT. sieboldii, in this area were 24.9％, 23.1％, mography of woody seedlings at cool-temperate natural for- and 26.5％, respectively7）. Although the dwarf bamboo est in central Japan. Browsing pressure from Sika deer (Cer- (Sasamorpha borealis (Hack.) Nakai) was widely distributed vus Nippon Temminck) has increased since, at least, the last in this region5）, retrogression occurred associated with deer two decades5，11，20）. Using a dominant tree species, Fagus ja- browsing at our study site (M. Kaji, pers. obs.). In 2006, no ponica Maxim., we monitored the date of seedling germina- dwarf bamboo but several shrub and herbal species (such as tion and its subsequent fate inside and outside deer exclo- Pieris japonica (Thunb.) D. Don ex G. Don, Pertya glabres- sure. We tested the advantage of early germination consider- cens Sch. Bip. ex Nakai, Ainsliaea acerifolia Sch. Bip. var. ing the effect of the presence of deer. We asked the follow- subapoda Nakai, and Chloranthus serratus Thunb.) can be ing questions. What was the limiting factor for germinated seen on the forest floor. Only P. japonica was an evergreen seedlings and what was its effect on seedling survival？ and unpalatable species for deer, which was the first major Was seedling demography, especially the advantage of early plants among the vegetation of the forest floor, despite its germination, affected by deer browsing？ patch distribution. In the growth season (from late spring to autumn), forage biomass increased with the emergence of 2. Materials and Methods the saplings of the upper tree species. Based on the periodic 2.1 Study species deer route-census survey by the Univ. Tokyo Forest, the av- Japanese beech, F. japonica, is a deciduous broad-leaved erage number of observed deer around our study site was tree. It is mainly distributed in the cool-temperate forests of 1.0／km in 2006 and varied with season; nearly 0.0／km during Pacific Ocean-type regions. In central Japan, F. japonica is December to February15）. 234 ISHIZUKA・KAJI・GOTO In this area, a mast year for F. japonica occurred during cotyle (or root in some cases) of the seedling was cut off, 2005 and a number of seedlings were expected to emerge in and／or that the apical meristem of the seedling was cut by 2006. In early April 2006, before the emergence of F. japon- herbivory. The former case was presumably attributed to ro- ica seedlings, we established a fenced exclosure to prevent dent attack, whereas the latter case was attributed to insect the effect of deer. We used 1.8 m height fencing poles made herbivory, such as larvae. Seedlings killed by fatal disease from steel pipes and polyethylene net to make a 28 × 14 m caused by pathogenic infections were classified as“damping plot in the fenced deer exclosure (i.e., the fenced plot). The off,”i.e., necrosis in the axis. Seedlings killed by drought or fenced plot was established immediately adjacent to a 50 × desiccation during the early root elongation phase were clas- 50 m plot, which was established in 1991 and divided into 25 sified as“desiccation.”The number of seedlings killed by subplots of 10 × 10marrangedin5× 5matrixes7）.Thelat- each mortality factor was calculated daily on the basis of the ter plot was outside the deer fence (i.e., the open plot).
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