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1 (Japanese Journal of Conservation Ecology) 15 : 3-15 (2010) Changes in the understory flora following Sika deer browsing with special reference to flowering stage in Makura-dani, Ashiu Exeperimental Forest Shinji Fujii University of Human Environments Abstract: Changes in the flora and the individual number of flowering plants under Sika deer browsing were investigated at Makura-dani in Ashiu Experimental Forest, Kyoto Prefecture, Japan. The number of species decreased from 84 to 56. Among the 77 species evaluated, eight species increased and 47 species decreased conspiciously in number, probably resulting in the local extinction of 22 species. The rate of species loss was greater for large-sized plants and the rate of unchanged or less changed species was greater for small-sized plants. Summer or autumn flowering herbaceous species showed higher rates of decreased species number than spring flowering species. Therefore, plant size and flowering season were the important characteristics for evaluating the browsing effect. The estimated number of flowering individuals and the Simpson's diversity index showed distinctly different phenological patterns after Shika deer browsing, and the seasonal function of flowering plant community on herbivorous insects visiting flowers were supposed to have changed. Keywords: flora change, number of flowering plants, phenology, plant size, plant community Cervus nippon Temminck Hypolepis punctata (Thunb.) Mett. ex Kuhn 2008 Kato and Okuyama University of Human Environments, Kami-sanbonmatsu 6-2, Motojyuku-cho, Okazaki-shi , Japan shinji@uhe.ac.jp

2 Kato and Okuyama Kato and Okuyama Maesa japonica (Thunb.) Moritzi ex Zoll Dryopteris expansa (C.Presl) Fraser-Jenk. et Jermy Veratrum album L. subsp. oxysepalum (Turcz.) Hultén Ligularia dentata (A.Gray) H.Hara 2000 Harper Kato and Okuyama 調査場所調査日時調査対象 1974 Mitella acerina Makino Cirsium ashiuense S.Yokoy. et T.Shimizu Yasuda and Nagamasu 1995 Yokoyama et al Arisaema amurense Maxim. subsp. robustrum (Engler) Ohashi et J.Murata var. ovale (Nakai) Ohashi et J.Murata Strobilanthes wakasana Wakas. et Naruh. Yasuda and Nagamasu 1995 Wakasugi and Naruhashi 1993 Hemerocallis dumortieri C.Morren var. esculenta (Koidz.) Kitam. ex M.Matsuoka et M.Hotta 1995 Gentiana triflora Pall. var. japonica (Kusn.) H.Hara Trautvetteria caroliniensis (Walter) Vail var. japonica (Siebold et Zucc.) T.Shimizu 1994 Yasuda and Nagamasu

3 1 1 1A Quercus crispula Blume Quercus serrata Thunb. ex Murray Fagus crenata Blume 1B Pterocarya rhoifolia Siebold et Zucc. Aesculus turbinata Blume 600 m 776 m 1,300 m 1A B 5 m 1,300 m 10 m 800 m 1A 500 m 1B 1980 Cryptomeria japonica (L.f.) D.Don Carpinus spp. Acer pictum Thunb. subsp. mayrii (Schwer.) H.Ohashi Acer micranthum Siebold et Zucc. Benthamidia japonica (Siebold et Zucc.) H.Hara Clethra barbinervis Siebold et Zucc. Stylax obassia Siebold et Zucc. Cerasus incisa (Thunb.) Loisel. var. kinkiensis (Koidz.) H.Ohba Menziesia cilicalyx (Miq.) Maxim. Sasa kurilensis (Rupr.) Makino et Shibata Elatostema japonicum Wedd. var. majus (Maxim.) H.Nakai et H.Ohashi Aconitum sanyoense Nakai Euonymus alatus (Thunb.) Siebold et Zucc. var. microphyllus (Nakai) Nakai Viola vaginata Maxim. Angelica polymorpha Maxim. Symplocos sawafutagi Nagam. Isodon trichocarpa (Maxim.) Kudo Carex persistens Ohwi Hydrocotyle spp. Centipeda minima (L.) A.Braun et Asch. データの収集と数値化

4 Elatostema laetevirens Makino N データの解析 /2 1/ cm 20 cm 20 cm Fisher 1975 景観変化 2 Persicaria thunbergii (Siebold et Zucc.) H.Gross Aster glehni Fr.Schm. var. hondoensis Kitam. Juncus effusus L. var. decipiens Buchenau Erechtites hieraciifolius (L.) Raf. ex DC 増減の種数と分類群 Potentilla centigrana Maxim. Daphne miyabeana Makino Pterostylax hispida Siebold et Zucc. Mimulus inflatus (Miq.) Nakai 2 6

シカ食害にともなう開花植物相の変化図 2 調査地の景観変化の例 a 1991 年 6 月 14 日のゴマナとキンキカサスゲが優占する植生景観 4 枚のパネルは本調査とは無関係 b a と同じ場所の 2007 年 6 月 16 日の植生景観イグサの優占群落に変化していた左奥のシカ防護ネットは本調査とは無関係 c 1991 年 6 月 27 日の植生景観アシウアザミが優占し

5 シカ食害にともなう開花植物相の変化図 2 調査地の景観変化の例 a 1991 年 6 月 14 日のゴマナとキンキカサスゲが優占する植生景観 4 枚のパネルは本調査とは無関係 b a と同じ場所の 2007 年 6 月 16 日の植生景観イグサの優占群落に変化していた左奥のシカ防護ネットは本調査とは無関係 c 1991 年 6 月 27 日の植生景観アシウアザミが優占し低木にはヤナギ類とヤブデマリがみられた d c と同じ場所の 2007 年 6 月 29 日の植生景観半裸地化しイグサとダンドボロギクの群落に変化していた中央のシカ防護ネットは本調査とは無関係 Cladothamnus bracteatus (Maxim.) T.Yamaz. ウスギヨウ表 1 増減が判明した 77 種の生育型と種数増加増加種数変化せず減少減少合計一年生草本多年生草本木本合計ラク Menziesia cilicalyx (Miq.) Maxim. アキチョウジ Isodon longitubus (Miq.) Kudo クロバナヒキオコシ I. trichocarpa (Maxim.) Kudo アキギリ Salvia glabrescens (Franch. et Sav.) Makino モミジガサ Cacalia delphiniifolia Siebold et Zucc. クルマバハグマ Pertya rigidula (Miq.) Makino の 10 分類群であったまた表 2 において * を内の数字は年調査時に 50 株以上の開花個体が確認された種数を示す付した 22 種は年の調査時に開花を確認していたが 2007 年の調査では開花個体と非開花個体の両方を発見することができなかった減少が認められた分類群のうち年の開花増減を評価できなかった 25 種の内訳は株数 1000 株以上から 2007 年の 10 株未満に激減したの年当時に生育していたかどうかの記憶が曖昧であるものはリョウメンシダ Arachniodes standishii (Moore) Ohwi 8 種年の開花を確認していたが当時の開ウワバミソウミゾソバの 3 種株から 10 花株数が不明なもの 14 種腐生植物のために毎年の出株未満に大きく減少したのはツリフネソウ Impatiens 現が期待できないもの 3 種であったなお 1989 textori Miq. ゴマナアシウアザミキンキカサスゲの 1994 年の開花株数の不明な植物 14 種のうちアカソ 4 種株から 10 株未満に減少したのはヤナギ Boehmeria silvestrii (Pamp.) W.T.Wang ムカゴイラクサ属 Salix spp. シラネセンキュウヒカゲミツバ Laportea bulbifera (Siebold et Zucc.) Wedd. イタドリ Spuriopimpinella koreana (Y.Yabe) Kitag. ホツツジ Fallopia japonica (Houtt.) Ronse Decr. イヌツゲ Ilex 7

6 Daphne miyabeana Pterostylax hispida Centipeda minima Erechtites hieraciifolius Juncus effusus var. decipiens Arisaema amurense subsp. robustrum var. ovale Anemone flaccida O2955* Chrysosplenium grayanum Oxalis acetosella subsp. griffithii Schizocodon soldanelloides var. magnus Shortia uniflora Symplocos sawafutagi K309 O2699 O3268 O2856 Potentilla centigrana O3341 Persicaria debilis O691 Mimulus inflatus N Aconitum sanyoense O Asarum nipponicum Corydalis lineariloba Chrysosplenium fauriae Mitella acerina M. furusei var. subramosa Euonymus sieboldianus Benthamidia japonica Gaultheria adenothrix Rhododendron molle subsp. japonicum Stylax obassia Ligustrum tschonoskii Veronica muratae O2861 O2949 K2968 K2952 N O2858 O2751 O2874 O2875 O3109 N O3030 Viburnum pilcatum var. O3034 tomentosum Arisaema serratum * OK N 8

7 ** Astilbe thunbergii * Aruncus dioicus var. tenuifolius * Salix spp. * Elatostema japonicum var. majus N O685 * Isodon Trichocarpa O3047 ** Salvia glabrescens O3145 O3152 Persicaria thunbergii O3146 * Aster glehni var. hondoensis O Impatiens textori O3149 ** Cacalia delphiniifolia N * Angelica polymorpha O3185 Cirsium ashiuense O3148 * Spuriopimpinella koreana O2864 ** Pertya rigidula O2854 ** Cladothamnus bracteatus O2860 ** Carex persistens O3042 ** Menziesia cilicalyx O2889 * Arachniodes standishii N ** Isodon longitubus O3155 Elatostema laetevirens O3050 * Chelonopsis moschata O * Silene miqueliana ** Stellaria monosperma var. japonica * Cardiandra alternifolia * Cerasus incisa * Ilex leucoclada * Euonymus alatus var. microphyllus N * Physaliastrum echinatum O4066 O3158 * P. savatieri O680 Melampyrum laxum O3046 var. nikkoense O2947 * Cacalia nikomontana O2855 Solidago virgaurea subsp. asiatica O2857 ** Synurus palmatopinnatifidus O3162 O3160 ** E. lanceolatus O2892 ** Cardiocrinum cordatum O3117 * E. oxyphyllus * Eleutherococcus divaricatus O4520 * Helonias orientalis N O680 O2958 O3792 * Paris tetraphylla N ** Panax japonicus O3051 Tricyrtis affinis ** Angelica pubescens O3110 Trillium apetalon O2951 ** Sanicula chinensis O3048 * Aniselytron treutleri var. japonicum ** Symplocos coreana O3343 Miscanthus tinctorius O3184 *2007 **2007 O682 9

8 3 57 <<20 cm 20 cm>>20 cm crenata Thunb. Sambucus racemosa L. subsp. sieboldiana (Miq.) H.Hara Aletris luteoviridis (Maxim.) Franch 生活形質と増減の傾向 P P P <<20 cm cm >>20 cm P P P P 0.01 P 0.24 推定開花株数の変化 >>20 cm 10

9 植生変化の要因 Platypus quercivorus 4 a b

10 (Murayama) 2 Hydrangea hirta (Thunb.) Siebold et Zucc Persicaria debilis (Meisn.) H.Gross ex W.T.Lee Panax japonicus C.A.Meyer Hydrangea serrata (Thunb.) Ser. 2.5 m Kato and Okuyama Kato and Okuyama 植物相と植生の変化 Crawley a, c b, d 2007 Arisaema serratum (Thunb.) Schott 12

11 植物体サイズ生活史型と増減 Kato and Okuyama M. furusei Ohwi var. subramosa Wakab. Cacalia nikomontana Matsum. Pertya rigidula (Miq.) Makino Cardiocrinum cordatum (Thunb.) Makino Trillium apetalon Makino 8 摂食の回避 Rhododendron molle (Blume) G.Don subsp. japonicum (A.Gray) K.Kron 2007 Anemone flaccida Fr.Schm. Corydalis lineariloba Siebold et Zucc. Oxalis acetosella L. subsp. griffithii (Edgew. et Hook.f.) H.Hara 5 Asarum nipponicum F.Maek. Schizocodon soldanelloides Siebold et Zucc. var. magnus (Makino) H. Hara Shortia uniflora (Maxim.) Maxim. 3 Kato and Okuyama Harper

12 5 群集の開花フェノロジーへの影響 Simpson Simpson Simpson Simpson Kato and Okuyama Bombus diversus deiversus Smith Bombus ardens ardens Smith Eobiana sp. Ognevia longipennis (Shiraki) Parapodisma sp. Stethophyma magister (Rehn) 2007 Kato and Okuyama

13 Crawley MJ (1997) Plant-herbivore dynamics. In: Crawley MJ (ed) Plant ecology 2nd edition. Blackwell Science, Oxford, pp (1996) 14. Nature Study 42(6):9-10 *Harper JL (1969) The role of predation in vegetational diversity. Brookhaven Symp Biol 22:48-62 Harper JL (1977) Population biology of plants. Academic Press, London (2000). 48:47-57 (1974)., (2008). 13: (1975)., (2006). ( ).,, pp (2006). Ecosophia 18:4-11 Kato M, Okuyama Y (2004) Changes in the biodiversity of a deciduous forest ecosystem caused by an increase in the Shika deer population at Ashiu, Japan. Contrib Biol Lab Kyoto Univ 29: (2001). 23:21-25 (1995). 46: (2007) (Cervus nippon). 24: (1989). 39:67-80 (2005). 10:11-17 (2007) (Cervus nippon) (Maesa japonica). 29:65-66 (2007). 12:20-27 (1992). 42: (1984). 7:1-6 Wakasugi T, Naruhashi N (1993) A new species, Strobilanthes wakasana (Acanthaceae) from Japan. J Phytogeogr Taxon 41:1-6 (2008)., (2002). ().,, pp (2008). ().,, pp (2001). 23: (1994). 45:77-78 Yasuda S, Nagamasu H (1995) Flora of Ashiu, Japan. Contrib Biol Lab Kyoto Univ 28: (2006). ( ).,, pp (2002) :15-18 Yokoyama S, Yamashita M, Shimizu T (1996) Taxonomic studies on the genus Cirsium (Compositae) in the Hokuriku district and adjacent areas (2) Cirsium ashiuense - a new species from Ashiu, Kyoto Prefecture. J Phytogeogr Taxon 44:19-24 (1993). ( ) 4.,, pp (2006)., * (1989) 15

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untitled 2006 50 2008 119 69 BG Plants YList 2001 Illicium anisatum L. Magnolia hypoleuca Siebold et Zucc. Magnolia kobus DC. Lindera praecox (Siebold et Zucc.) Blume Lindera umbellata Thunb. Machilus thunbergii