植物科学最前線 5:93 （2014）⳦ᚑᒓᰤ㣴ᛶࡢ⏕άᵝᘧࢆྍ⬟࡟ࡋࡓᵝࠎ࡞㐺ᛂ㐍໬̿≉࡟㏦⢊ᵝᘧࡢኚ໬࡟ࡘ࠸࡚
䢢䢢ᮎḟ೺ྖ࣭ຍ⸨┿ி㒔኱Ꮫ኱Ꮫ㝔ே㛫࣭⎔ቃᏛ◊✲⛉ࠛி㒔ᕷᕥி༊ྜྷ⏣஧ᮏᯇ⏫Kenji Suetsugu, Makoto Kato
Evolution of life history traits suited to mycoheterotrophic life stylewith special reference to pollination biology
Key words: breeding system, complex adaptive trait, mixotrophy, mycoheterotrophy,
Orchidaceae, reproductive biology, self-pollination
Graduate School of Human and Environmental Studies, Kyoto University
Yoshida Nihonmatsu-cho, Sakyo, Kyoto 606-8501, Japan㸯ࡣࡌࡵ࡟
ከࡃࡢ㝣ୖ᳜≀ࡣ᰿ᅪ࡟࠾࠸࡚⳦᰿⳦࡜ඹ⏕㛵ಀࢆ⤖ࢇ࡛࠾ࡾ㸪᳜≀ࡣගྜᡂ࡛ᚓࡓ
ྠ໬⏘≀ࢆ⳦᰿⳦࡟ᥦ౪ࡋ㸪ࡑࡢぢ㏉ࡾ࡟⳦᰿⳦ࡣỈࡸ❅⣲ࡸࣜࣥ࡞࡝ࡢ↓ᶵሷ㢮ࡢ྾཰ࡸ㜵⾨ࡢᡭຓࡅࢆࡋ࡚࠸ࡿࠋ
ࡋ࠿ࡋ࡞ࡀࡽ㸪᳜≀ࡢ୰࡟ࡣ㸪ගྜᡂ⬟ຊࢆኻ࠸㸪௚ࡢ⏕≀࠿ࡽࡍ࡭࡚ࡢ㣴ศࢆ␎ዣࡍࡿ࡜࠸
࠺≉␗࡞㐍໬ࢆ㐙ࡆࡓᚑᒓᰤ㣴᳜≀ࡀᏑᅾࡍࡿࠋࡇࢀࡽࡢ᳜≀ࡣ㸪௚ࡢ᳜≀࡟ྲྀࡾ௜࠸࡚㣴ศࢆዣ࠺ᐤ⏕᳜≀࡜㸦Suetsugu etal. 2008, 2012a㸧㸪⳦᰿⳦࠿ࡽ㣴ศࢆዣ࠺⳦ᚑᒓᰤ㣴᳜≀࡟኱ูࡉࢀࡿ 㸦Leake 1994, Bidartondo
2005; ᅗ 1㸧ࠋ≉࡟㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪㛤ⰼ㸪⤖ᐇᮇ௨እࡣᆅୖ࡟ጼࢆ⌧ࡉࡎ㸪᳜≀య⮬యࡀぢࡘ࠿ࡾ࡟ࡃ࠸ࡓࡵ㸪᳜≀┦ࡢ◊✲ࡀ㐍ࢇᅗ ᐤ⏕᳜≀࡜⳦ᚑᒓᰤ㣴᳜≀ ୧⪅࡜ࡶගྜᡂ⬟ࢆಖᣢࡋࡓࡶࡢ࡜᏶඲࡟႙ኻࡋࡓࡶࡢࡀ࠶ࡿࠋᐤ⏕᳜≀ࡣ྾ჾ࡛௚ࡢ᳜≀࡟ྲྀࡾ௜ࡃࡇ࡜࡛㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ⳦⣒ࢆ⣽⬊୰࡟ྲྀࡾ㎸ࡴࡇ࡜࡛㸪ᐤ⏕ࢆᡂ❧ࡉࡏࡿ 

K. Suetsugu - 1

植物科学最前線 5:94 （2014）ࡔ᪥ᮏ࡛ࡉ࠼㸪࡯࡜ࢇ࡝ࡢ✀㢮࡟࠾࠸࡚ṇ☜࡞ศᕸ᝟ሗࡣㅦࡢࡲࡲ࡛࠶ࡿ 㸦Suetsugu &Ishida 2011, Suetsugu et al. 2012b, Suetsugu 2013, Suetsugu et al. 2013 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ
⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᪤▱ࡢ✀ᩘࡇࡑࡑࢀ࡯࡝ከࡃ࡞࠸ࡶࡢࡢ㸪ᖜᗈ࠸ศ㢮⩌࡛ぢࡽࢀ㸪⿕Ꮚ᳜≀ࡔࡅ࡛ࡶ㸪ࢶࢶࢪ⛉㸪ࣄ࣓ࣁࢠ⛉㸪ࣜࣥࢻ࢘⛉㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘⛉㸪ࢥࣝࢩ࢔⛉㸪࢔࣓ࣖ⛉㸪ࣛࣥ⛉㸪ࢧࢡࣛ࢖ࢯ࢘⛉㸪࣍ࣥࢦ࢘ࢯ࢘⛉࡟࠾࠸࡚㸪ࡑࢀࡒࢀ⊂❧࡟㉳※ࡋ࡚࠸ࡿࡇ࡜ࡀࢃ࠿ࡗ࡚࠸ࡿ 㸦Leake 1994, Bidartondo 2005㸧ࠋࡇࢀࡲ࡛㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ወ␗࡞ᙧែ࠿ࡽேࠎࡢ㛵ᚰࢆ㞟ࡵ࡚ࡁࡓࡀ㸪Ꮫၥⓗ࡞᥈ồࡣ༑ศ࡛ࡣ࡞࠿
ࡗࡓࠋࡼࡗ࡚㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ࡝ࡢࡼ࠺࡟㉳※ࡋ㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿ࢆ᫂
ࡽ࠿࡟ࡍࡿࡇ࡜ࡣ㸪᳜≀Ꮫ࡟࠾ࡅࡿ㔜せ࡞ㄢ㢟࡛࠶ࡿ࡜ゝ࠼ࡿࡔࢁ࠺ࠋ ㏆ᖺ㸪ศᏊ⏕≀ᏛⓗᡭἲࡸᏳᐃྠ఩యศᯒ࡜࠸ࡗࡓᡭἲࡢⓎ㐩࡟క࠸㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ㐍໬ࡢ㐣⛬࡛㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋ౛࠼ࡤ㸪⥳ⴥࢆᣢࡘ✀㢮࡜᏶඲࡟ⴥ⥳⣲ࢆᣢࡓ࡞࠸✀ࢆྵࡴ㏆⦕✀ࡢ⣔⤫ᶞࢆసᡂࡍࡿࡇ࡜࡛㸪᏶඲࡟ⴥ⥳⣲ࢆኻࡗࡓ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࡢฟ⌧࡟ඛ⾜ࡋ࡚㸪⥳ⴥࢆᣢࡕ࡞ࡀࡽ㸪⮬㌟ࡢ
ගྜᡂ⏘≀ࡢ୙㊊ࢆ⿵࠺㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࡀඛ࡟⌧ࢀࡿሙྜࡀ࠶ࡿࡇ࡜ࡀᣦ᦬
ࡉࢀ࡚࠸ࡿ 㸦Selosse and Roy 2009; ➨㸰❶㸪➨㸴❶ཧ↷㸧ࠋࡲࡓࡑࡢ⣔⤫ᶞୖ࡟㸪⳦࡬ࡢ౫Ꮡᗘࡸඹ⏕⳦ྀ࡜࠸ࡗࡓᙧ㉁ࢆ࣐ࢵࣆࣥࢢࡍࡿࡇ࡜࡛㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡟క࠸㸪
ࡼࡾⅣ⣲※ࢆዣ࠸ࡸࡍ࠸⳦㢮࡬ࡢᐟ୺㌿᥮ࡸ≉Ṧ໬ࡀ㉳ࡇࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗ࡚ࡁ
ࡓ 㸦Selosse & Roy 2009㸧ࠋୖグࡢࡼ࠺࡟⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ㐍໬ࡢ㐣⛬࡛⳦࡜ࡢ㛵ࢃࡾࢆኚ໬ࡉࡏ࡚ࡁࡓࡇ
࡜ࡣ㸪㠀ᖖ࡟ࢃ࠿ࡾࡸࡍ࠸ࢫࢺ࣮࣮࡛ࣜ࠶ࡾ㸪ཷࡅධࢀࡸࡍ࠸ࡶࡢ࡛࠶ࡿࠋࡋ࠿ࡋ࡞ࡀ
ࡽ㸪ࠕ⳦ᚑᒓᰤ㣴᳜≀ࡀࡑࡢ≉␗࡞⏕άྐࢆ඲࠺ࡍࡿࡓࡵ࡟࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿ࠖࢆ⌮ゎࡍࡿࡓࡵ࡟ࡣ㸪ᆅୗ㒊࡜ᆅୖ㒊ࡢ୧᪉࡛㸪࡝ࡢࡼ࠺࡞㐺ᛂࡀぢࡽࢀࡿࡢ࠿ࢆ▱ࡿᚲせࡀ࠶ࡿࡣࡎ࡛࠶ࡿࠋᮏ✏࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆ㸪⏕Ꮡࢆྍ⬟࡜ࡋ࡚࠸ࡿࡢ࠿ࢆ㸪ࡇࢀࡲ࡛࠶ࡲࡾὀ┠ࡉࢀ࡚࠸࡞࠿ࡗࡓᆅୖ㒊࡛ࡢ㐺ᛂࢆྵࡵ⤂௓ࡋࡓ࠸ࠋ

㸰⳦ᚑᒓᰤ㣴ᛶࡢ㐺ᛂⓗព⩏ࡲࡎ⳦㢮࠿ࡽ㣴ศࢆ␎ዣࡍࡿ࡜࠸࠺㐍໬࡟㸪࡝ࡢࡼ࠺࡞㐺ᛂⓗព⩏ࡀ࠶ࡿࡢ࠿⪃࠼࡚ࡳࡼ࠺ࠋ୍⯡ⓗ࡟ᗈࡃཷࡅධࢀࡽࢀ࡚࠸ࡿㄝ᫂ࡣ㸪⳦࡟ᐤ⏕ࡍࡿࡇ࡜࡛㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛ࡶ⏕Ꮡྍ⬟࡟࡞ࡗࡓ࡜࠸࠺ࡶࡢ࡛࠶ࡿ 㸦Bidartondo et al. 2004㸧ࠋࡋ࠿ࡋ㸪ᬯ࠸ᯘᗋ࡛ࡢ⏕⫱ࢆྍ⬟࡟ࡍࡿࡇ࡜௨እ࡟ࡶ㸪⳦࡬ࡢ౫Ꮡᛶࢆ㧗ࡵࡿࡇ࡜ࡢ㐺ᛂⓗព⩏ࡀ࠶ࡿྍ⬟ᛶࡣᤞ࡚ࡁࢀ࡞࠸ࠋ ᡃࠎࡣ㸪⳦࡟ᐤ⏕ࡍࡿࡇ࡜ࡢูࡢ㐺ᛂⓗព⩏࡜ࡋ࡚㸪୍ᗘ㸪ᆅୖ㒊ࢆ⌧ࡋࡓᚋ࡛ࡶ㸪ࢫࢺࣞࢫ࡟ࡉࡽࡉࢀࡓ㝿࡟ࡣ㸪⩣ᖺ㸪ᆅୗ࡟࡜࡝ࡲࡾ㸪⳦࠿ࡽ༑ศ࡞Ⅳ⣲※ࢆᚓࡓᚋ㸪෌ᗘ㸪ᆅୖ㒊ࢆ⌧ࡍ࡜࠸࠺ࡇ࡜ࡀ࡛ࡁࡿ࡜࠸࠺฼Ⅼࢆ⪃࠼ࡓࠋ㏻ᖖࡢගྜᡂࡔࡅࢆ⾜ࡗ࡚࠸ࡿ᳜≀࡟࠾࠸࡚ࡶ㸪ከᖺ⏕᳜≀ࡢሙྜ㸪ᆅୖ࡟⌧ࢀ࡞࠸≧ែ࡛Ꮡᅾࡋ⥆ࡅ㸪⩣ᖺ௨

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植物科学最前線 5:95 （2014）㝆㸪෌ᗘ㸪ᆅୖ㒊ࡀ⌧ࢃࢀࡿࡇ࡜ࡀ࠶ࡿࠋࡇ
࠺ࡋࡓ⌧㇟ࡣࠕᆅୖჾᐁࡢఇ╀ 㸦vegetative
dormancy ࡢヂㄒ ௨ୗࡣ㸪༢࡟ఇ╀࡜ࡍࡿ㸧ࠖ࡜࿧ࡤࢀࡿࡀ 㸦Shefferson 2009㸧㸪✀Ꮚࡢఇ╀
࡜ࡣ␗࡞ࡾ㸪᰿ࡢ࿧྾࡞࡝ࡢ௦ㅰάືࡣ೵Ṇࡋ࡞࠸ࠋࡑࡢࡓࡵࡇࢀࡲ࡛࡟ㄪᰝࡉࢀࡓ࡯࡜
ࢇ࡝ࡢ᳜≀࡛㸪⵳✚ࡋࡓ㣴ศࡀᯤῬࡋ࡚ࡋࡲ
࠸㸪ఇ╀ࢆ⤒ࡓಶయࡢ⏕Ꮡ⋡ࡀపୗࡍࡿࡇ࡜
ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Gregg & Kéry 2006,
Hutchings 1987, Shefferson et al. 2003, Shefferson& Tali 2007 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ⳦࡟ᐤ⏕ࡍࡿ᳜≀ࡢሙྜ㸪ఇ╀୰ࡶ㸪⳦㢮࠿ࡽⅣ⣲※ࢆ⿵࠺ࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪ఇ╀ࡢࢥࢫࢺࡀ㍍ῶࡉ
ࢀ㸦ࡴࡋࢁᆅୖ㒊ࢆ⌧ࡉ࡞࠸ࡇ࡜࡛㣗ᐖ࡞࡝ࢆཷࡅࡿ༴㝤ᛶࡀῶࡿࡓࡵ㸧㸪⏕Ꮡ⋡ࢆಖࡘࡇᅗ 㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢟ࣥࣛࣥ

࡜ࡀ࡛ࡁࡿྍ⬟ᛶࡀ࠶ࡿࠋ ࡑࡇ࡛ᡃࠎࡣ㸪Mark and recapture ἲࢆ⏝࠸㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ࢟ࣥࣛࣥࡢఇ╀⋡ࡸ⏕Ꮡ⋡ࡀ࡝ࡢ⛬ᗘ࠿㸪ࡉࡽ࡟㣗ᐖࡸ⤖ᐇࡢ᭷↓࡞࡝࡟ࡼࡾ㸪ఇ╀⋡ࡸ⏕Ꮡ⋡ࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡍࡿࡢ࠿ࢆ᳨ウࡋࡓࠋ࢟ࣥࣛࣥࡣ᱂ᇵࡀ㞴ࡋࡃ㸪㛗ᮇ㛫⥔ᣢࡍࡿࡇ࡜ࡣ㞴ࡋ࠸ࡓࡵ㸪⥳ⴥࢆᒎ㛤ࡋ࡚࠿ࡽࡶ⳦࡟౫Ꮡࡋ࡚࠸ࡿྍ⬟ᛶࡀ♧၀ࡉࢀ࡚࠸ࡓࡀ㸪㏆ᖺ⾜ࢃࢀࡓ᱂ᇵᐇ㦂ࡸᏳᐃྠ఩యศᯒ࠿ࡽ㸪ᐇ㝿࡟⳦࡟౫Ꮡࡋ࡚࠸ࡿࡇ࡜ࡀド᫂ࡉࢀ࡚࠸ࡿ᳜≀࡛࠶ࡿ 㸦ᅗ 2; Yagame & Yamato 2013, ᆏᮏࡽ 2013㸧ࠋࡑࡢ⤖ᯝ㸪ᡃࠎࡢண᝿㏻ࡾ㸪㣗ᐖࢆཷࡅࡓ࡜ࡁࡸ⤖ᐇࡋࡓ࡜ࡁ࡞࡝㸪᳜≀ࡢ㈨※ࡀᯤῬࡋࡓ㝿࡟ࡣ㸪ఇ╀ࡀ㉳ࡇࡾࡸࡍࡃ࡞ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࡲࡓࡇࡢࡼ࠺࡟⏕Ꮡࡀ㞴ࡋࡃ࡞ࡗࡓ≧ែ࡛ఇ╀࡟ධࡿ࡟ࡶ㛵ࢃࡽࡎ㸪ఇ╀ࢆ⤒ࡓಶయࡢ⏕Ꮡ⋡ࡀపୗࡍࡿࡇ࡜ࡣ࡞࠿ࡗࡓ 㸦ᮎḟࡽ ᢞ✏୰㸧ࠋᐇࡣ㸪ඛ⾜◊✲࡛ࡶ㸪౛እⓗ࡟ఇ╀ᚋࡢ⏕Ꮡ⋡ࡀୗࡀࡽ࡞࠸✀㢮ࡀ▱ࡽࢀ࡚࠸ࡿࠋࡇ
ࢀࡽࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢほⅬ࠿ࡽࡣ㆟ㄽࡉࢀ࡚࠸࡞࠿ࡗࡓࡀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ✀㢮࡛」ᩘᅇ㸪ሗ࿌ࡉࢀ࡚࠸ࡿ⌧㇟࡛࠶ࡿ 㸦Shefferson et al. 2005, Jäkäläniemi et al. 2011㸧ࠋୖグࡢࡇ࡜ࢆ⪃࠼ేࡏࡿ࡜㸪ࠕࢫࢺࣞࢫࢆཷࡅࡓ㝿࡟㸪ᆅୖ㒊ࢆᒎ㛤ࡏࡎ࡟ᆅୗ࡟࡜࡝
ࡲࡿࡇ࡜ࡀ࡛ࡁࡿࠖࡇ࡜ࡶ㸪⳦࡟ᐤ⏕ࡍࡿព⩏ࡢ୍ࡘ࡛࠶ࡿ࡜⪃࠼ࡿࡇ࡜ࡀ࡛ࡁࡿࠋ
䢢㸱࢔ࣝࣅࣀ✺↛ኚ␗య࠿ࡽぢ࠼ࡿ↓ⴥ⥳໬ࡢ㞀ቨ
ඛ࡟㏙࡭ࡓࡼ࠺࡟㸪ࣛࣥ⛉ࡸࢶࢶࢪ⛉࡞࡝ࡢ୍㒊ࡢ✀㢮ࡣ㸪Ⓨ㐩ࡋࡓ⥳ⴥࢆᒎ㛤ࡋ㸪୍ぢࡍࡿ࡜ගྜᡂࡔࡅ࡛⏕Ꮡྍ⬟ࡢࡼ࠺࡟ぢ࠼ࡿ࡟ࡶ࠿࠿ࢃࡽࡎ㸪⳦㢮࡟ࡶⅣ⣲ࢆ୍㒊౫Ꮡࡋ࡚࠸ࡿ✀㢮ࡀᏑᅾࡍࡿ 㸦Gebauer & Meyer 2003, Bidartondo et al. 2004, Tedersoo et
al. 2007 ࡞࡝㸧ࠋࡇࡢࡼ࠺࡞㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࢆྵࡴࢢ࣮ࣝࣉ࠿ࡽ㸪ⴥ⥳⣲ࢆኻࡗࡓ

K. Suetsugu - 3

植物科学最前線 5:96 （2014）᏶඲࡟⳦ᚑᒓᰤ㣴ᛶ࡟≉໬ࡋࡓ✀㢮ࡀฟ⌧ࡍࡿࡇ࡜ࡀ࠶ࡿࡀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢ✀ᩘ࡟ẚ࡭࡚㸪᏶඲࡟⳦ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋࡓ✀ᩘࡣࡁࢃࡵ࡚ᑡ࡞࠸ࠋ๓㡯࡛㏙࡭ࡓࡼ࠺࡟
⳦࡟ᐤ⏕ࡍࡿ࡜࠸࠺⏕άྐࡀ᳜≀࡟࡜ࡗ࡚ᚓ⟇࡛࠶ࡿ࡞ࡽࡤ㸪⳦࠿ࡽࡢ㣴ศ␎ዣࡔࡅ࡛
⏕⫱ࡍࡿ↓ⴥ⥳᳜≀ࡣఱᨾᑡ࡞࠸ࡢࡔࢁ࠺࠿ࠋࡇࡢࡇ࡜ࢆ⪃࠼ࡿࡓࡵ࡟㸪↓ⴥ⥳໬ࡢ⋓ᚓ࡟࡝ࡢࡼ࠺࡞㞀ቨࡀ࠶ࡿࡢ࠿ࢆ⪃࠼࡚ࡳࡼ࠺ࠋ ⯆࿡῝࠸ࡇ࡜࡟㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᏶඲࡟ⴥ⥳⣲ࢆኻࡗࡓ࢔ࣝࣅࣀ✺↛ኚ␗యࢆ⏕ࡌࡿࡇ࡜ࡀከ࠸ 㸦Selosse et al. 2004, Julou etal. 2005, Tranchida-Lombardo et al. 2010 ࡞࡝㸧ࠋ
㏻ᖖࡢ᳜≀࡛࠶ࢀࡤ㸪ⴥ⥳⣲ࢆኻࡗࡓ࢔ࣝࣅ
ࣀࡣ㸪✀Ꮚ࡟㈓ⶶࡉࢀࡓ㣴ศࢆ౑࠸ᯝࡓࡍ࡜
ᯤࢀ࡚ࡋࡲ࠺ࠋࡋ࠿ࡋ࡞ࡀࡽ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀࡣ㸪⳦࠿ࡽⅣ⣲※ࢆྵࡴ㣴ศࢆ␎ዣࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪ⴥ⥳⣲ࢆᣢ
ࡘ㏻ᖖಶయ࡜ྠ⛬ᗘࡲ࡛ᡂ㛗ࡋ㸪ⰼࢆဏ࠿ࡏ
ࡿࡇ࡜ࡀ࡛ࡁࡿࡢ࡛࠶ࡿࠋࡇࡢࡇ࡜ࡣ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪⳦࠿ࡽ␎ዣࡋࡓ㣴ศࡔࡅ࡛⏕Ꮡࡍࡿࡇ࡜ࡀྍ⬟࡞ࡇ࡜ࢆ♧၀ࡋ࡚࠸ࡿࠋ
࡟ࡶ࠿࠿ࢃࡽࡎ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀࡣ⛥࡞Ꮡᅾ࡛࠶ࡾ㸪㞟ᅋෆ࡟ᗈࡀࡿࡇ
࡜ࡣࡵࡗࡓ࡟࡞࠸ࡢ࡛࠶ࡿࠋᅗ ࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀ✺↛ኚ␗య⳦࠿ࡽ㣴ศࢆ␎ዣࡍࡿ⬟ຊࢆ⋓ᚓࡋࡓࢢ࣮ࣝࣉ
ࡢ୰࡛㸪᏶඲࡟↓ⴥ⥳࡟࡞ࡗࡓ✀㢮ࡀᑡ࡞࠸ࡇ࡜ࡸ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀ
ࡀ㸪㞟ᅋෆ࡟ᗈࡀࡽ࡞࠸ࡇ࡜ࡣ㸪⳦࡟ᑐࡍࡿᐤ⏕⬟ຊࡢ⋓ᚓ௨እࡢ㠃࡛ࡶ㸪↓ⴥ⥳໬ࢆ㐩ᡂࡍࡿࡓࡵ࡟ࡣࡉࡽ࡞ࡿ㐺ᛂࡀᚲせ࡛࠶ࡿࡇ࡜ࢆ♧၀ࡋ࡚࠸ࡿࠋ ࢟ࣥࣛࣥࡶ௚ࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ౛࡟ࡶࢀࡎ㸪ࡋࡤࡋࡤ࢔ࣝࣅࣀࢆ⏕ࡌࡿಶయ⩌ࡀぢཷࡅࡽࢀࡿ 㸦ᅗ 3㸧ࠋࡇࡢࡼ࠺࡞࢔ࣝࣅࣀࡀఱᨾ㞟ᅋ୰࡟ᗈࡀࡗ࡚࠸࠿࡞࠸ࡢ࠿
㸦ࡘࡲࡾ㸪↓ⴥ⥳ࡢ㐍໬࡟ࡣ࡝ࡢࡼ࠺࡞㞀ቨࡀ࠶ࡿࡢ࠿㸧 ࢆ㸪࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀ࡜
㏻ᖖಶయࡢ✀Ꮚ⏕⏘ຊࢆ㐺ᛂᗘࡢᣦᶆ࡜ࡍࡿࡇ࡜࡛⪃࠼࡚ࡳࡓ࠸ࠋ࢟ࣥࣛࣥࡣ㸪㏦⢊⪅ࡀ࠸࡞ࡅࢀࡤᯝᐇࢆࡘࡅࡿࡇ࡜ࡀฟ᮶࡞࠸᳜≀࡛㸪ࣄ࣓ࣁࢼࣂࢳࡸࢥࣁࢼࣂࢳ࡜࠸ࡗࡓ
ᑠᆺࡢࣁࢳ࡟ⰼ⢊ࡢ፹௓ࢆクࡋ࡚࠸ࡿ 㸦ᮎḟࡽ ᢞ✏୰㸧ࠋࡼࡗ࡚㸪࢔ࣝࣅࣀࡢ࡯࠺ࡀ
㏦⢊⪅ࡢㄏᘬຊࡀஈࡋࡅࢀࡤ㸪ᯝᐇࢆࡘࡅࡿᩘ㸪ࡦ࠸࡚ࡣ✀Ꮚᩘ࡟㐪࠸ࡀฟࡿ࡜⪃࠼ࡽ
ࢀࡿࠋ㏦⢊⪅ࡢゼࢀࡓᅇᩘࢆṇ☜࡟ᢕᥱࡍࡿࡇ࡜ࡣ㸪㏻ᖖࡢ᳜≀࡟࠾࠸࡚ࡣ㛤ⰼᮇ㛫୰⥅⥆ࡋ࡚ほᐹࡋ࡞࠸㝈ࡾᴟࡵ࡚㞴ࡋ࠸ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࣖࢡࢩ࣐ࣛࣥள⛉ࢆ㝖ࡃࣛࣥ⛉࡛ࡣ㸪ほᐹࡀᐜ࡛᫆࠶ࡿ 㸦Suetsugu & Fukushima 2013, Suetsugu & Tanaka 2013a, b㸧ࠋఱᨾ࡞ࡽࡤ㸪ᩘ༑୓ࡢⰼ⢊ࡀ஫࠸࡟⤖ྜࡋ࡚ሢ࡟࡞ࡗࡓⰼ⢊ሢ࡜࿧ࡤࢀࡿᵓ㐀࡜࡞ࡗ࡚࠾

K. Suetsugu - 4

植物科学最前線 5:97 （2014）ࡾ㸪ࡑࡢࡓࡵ඲࡚ࡢⰼ⢊ࡣ㸪᪻⹸ࡀ 1 ᅇ㸦ከࡃ࡚ࡶᩘᅇ㸧ゼࢀࡓࡔࡅ࡛ᣢࡕཤࡽࢀࡿ࠿
ࡽ࡛࠶ࡿࠋࡑࡇ࡛ᐇ㝿࡟ⰼ⢊ࡢᣢࡕཤࡾ⋡ࡸཷ⢊⋡ࢆㄪ࡭ࡓ࡜ࡇࢁ㸪࢔ࣝࣅࣀಶయࡶ㏻
ᖖಶయ࡜ྠ⛬ᗘ㸪㏦⢊⪅ࢆ࠾ࡧࡁᐤࡏࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ㸪࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀಶయࡢ㐺ᛂᗘࡀୗࡀࡿ⌮⏤ࡣ㸪㏦⢊⪅ࡢㄏᘬຊࡢ
ᕪ࡛ࡣ࡞࠸ࡢ࡛࠶ࡿࠋࡇࡢᚋࡢ㡯┠࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ཷ⢊⏕ែࡢ㠃࡛ࡶไ⣙ࢆཷࡅ࡚࠸ࡿࡇ࡜ࢆ㏙࡭ࡿࡀ㸪ࡇࢀ࡟ࡣᬯᡤ࡬ࡢ㐍ฟࡀ῝ࡃ㛵ࢃࡗ࡚࠸ࡿࠋࡋ࠿ࡋ࡞ࡀࡽ
࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀࡣ㸪㏻ᖖಶయ࡜ྠࡌࡼ࠺࡞ග⎔ቃ࡟⏕⫱ࡋ࡚࠸ࡿࡓࡵ㸪ཷ⢊ࡢ㠃࡛㸪ⴭࡋ࠸୙฼┈ࢆ⿕ࡿࡇ࡜ࡣ࡞࠸ࡢ࡛࠶ࡿࠋ㐺ᛂᗘࡀୗࡀࡿ⌮⏤ࡀ㏦⢊⪅࡜ࡣ↓㛵ಀ࡛࠶ࡿࡇ࡜ࡣ㸪⮬ື⮬ᐙཷ⢊ࡀྍ⬟࡞ࢠࣥࣛࣥࡸࣁ࣐࡛࢝࢟ࣛࣥࡶ࢔ࣝࣅࣀಶయࡶ㞟ᅋෆ࡟ᗈࡀࡗ࡚࠸࡞࠸ࡇ࡜࠿ࡽࡶ♧၀ࡉࢀࡿ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋ ࡑࢀ࡛ࡣఱᨾ࢔ࣝࣅࣀࡣ㞟ᅋෆ࡟ᗈࡀࡽ࡞࠸ࡢ࡛࠶ࢁ࠺࠿ࠋ࢔ࣝࣅࣀ࡜㏻ᖖಶయࡢ㑇ఏⓗ࡞Ỵᐃ࣓࢝ࢽࢬ࣒ࡣ᫂ࡽ࠿࡛ࡣ࡞࠸ࡶࡢࡢ㸪࢔ࣝࣅࣀࡀຎໃ࡛࣍ࣔ࠶ࡿྍ⬟ᛶࡣ༑ศ⪃࠼ࡽࢀ㸪ࡇࡢࡇ࡜ࡀ୍ᅉ࡜࡞ࡗ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋࡋ࠿ࡋ࡞ࡀࡽほᐹࢆ⥆ࡅ࡚࠸ࡿ࡜㸪ࡑࡢ௚ࡢ⌮⏤ࡢᏑᅾࡶ♧၀ࡉࢀࡓࠋ୰࡛ࡶ኱ࡁ࡞⌮⏤࡜ࡋ࡚࢔ࣝࣅࣀࡣ㏻ᖖಶ
యࡼࡾࡶ᪩࠸ࢱ࢖࣑ࣥࢢ 㸦ከࡃࡢሙྜ✀Ꮚࡀᡂ⇍ࡍࡿ๓㸧 ࡟ᾘኻࡋ࡚ࡋࡲ࠺ࡓࡵ㸪㏻
ᖖಶయ࡟ẚ࡭ 1%௨ୗࡢ✀Ꮚࡋ࠿ṧࡍࡇ࡜ࡀ࡛ࡁ࡞࠸ࡇ࡜ࡀᣲࡆࡽࢀࡿࠋ⤖ᐇ๓࡟ᯤࢀ
࡚ࡋࡲ࠺ࡢࡣ㸪ගྜᡂࢆ⾜ࢃ࡞࠸࡟ࡶ㛵ࢃࡽࡎ኱ࡁ࡞ⴥࡸẼᏍࢆಖᣢࡋ࡚࠸ࡿࡓࡵ㸪⵨ᩓࡀ࠺ࡲࡃ࠸࠿ࡎ஝⇱ࡋࡸࡍࡃ㏻ᖖಶయࡼࡾ᪩ࡃᯤࢀ࡚ࡋࡲ࠺ࡇ࡜ࡀ୍ᅉࡔ࡜⪃࠼ࡽࢀࡿ 㸦Roy et al. 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓ࢔ࣝࣅࣀࡣⴥ⥳⣲ࢆᣢࡘ㏻ᖖ
ಶయ࡜ྠ⛬ᗘࡲ࡛ᡂ㛗ࡍࡿࡶࡢࡢ㸪ගྜᡂ⬟ࢆኻࡗ࡚࠸ࡿศࡔࡅⅣ⣲ไ㝈࡟㝗ࡾࡸࡍ࠸ྍ⬟ᛶࡶ࠶ࡿࠋࡑࡢሙྜ㸪ྠ໬ჾᐁࡢ㏥໬ࡣ㸪⵨ᩓ㐣ከ࡟࡞ࡿࡢࢆ㜵ࡄࡔࡅ࡛࡞ࡃ㸪㈨※๐ῶࡢព࿡࡛ࡶ኱ࡁ࡞ព࿡ࢆᣢࡘ࡜⪃࠼ࡽࢀࡿࠋ௚࡟ࡶ㸪㸦1㸧 ࢔ࣝࣅࣀಶయࡣ㏻ᖖ
ಶయࡼࡾࡶ㸪⳦࡬ࡢ౫Ꮡᗘࡀ㧗࠸ࡓࡵ❅⣲ྵ᭷⋡ࡀ㧗ࡃ㸪᳜㣗ᛶ᪻⹸࡟࡜ࡗ࡚ዲ㐺࡞㣵࡟࡞ࡿࡇ࡜ࡸ㸪㸦2㸧 ཯ᑕ⋡ࡢ㧗࠸ⓑ࠸ⴥࢆᣢࡘࡇ࡜࡛㸪᳜㣗ᛶ᪻⹸࠿ࡽぢࡘࡅࡽࢀࡸ
ࡍࡃ࡞ࡿࡇ࡜࡟ࡼࡿ㣗ᐖ⋡ࡢୖ᪼࡞࡝ࡶ࢔ࣝࣅࣀࡢ⏕⫱࡟ࡣ୙฼࡟ാ࠸࡚࠸ࡿ
㸦Stöckel et al. 2011, Roy et al. 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ↓ⴥ⥳᳜≀࡟࡞ࡿࡓࡵ࡟ࡣ㸪⳦࠿ࡽࡢ㣴ศ␎ዣࢆྍ⬟࡟ࡍࡿ⬟ຊࡢࡳ࡞ࡽࡎ㸪ẼᏍࡢ㏥໬㸪ⴥ࡟௦⾲ࡉࢀࡿྠ໬ჾᐁࡢ㏥໬㸪㛤ⰼ࠿ࡽ⤖ᐇࡲ࡛ࡢᮇ㛫ࡢ▷⦰㸪㠀㛤ⰼ᫬ᮇࡢఇ╀㸪ಖㆤⰍ㸪⦾
Ṫᵝᘧࡢኚ໬ 㸦⦾Ṫᵝᘧࡢኚ໬࡟ࡘ࠸࡚ࡣḟ㡯࡛ヲࡋࡃㄝ᫂ࡍࡿ㸧 ࡞࡝ࡢᵝࠎ࡞㐺ᛂࢆ㐙ࡆࡿᚲせࡀ࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋࡇࢀࡽࡢ㐺ᛂࢆྠ᫬࡟㐙ࡆࡿࡇ࡜ࡀ㞴ࡋ࠸ࡓࡵ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡟ẚ࡭㸪ⴥ⥳⣲ࢆኻࡗࡓ⳦ᚑᒓᰤ㣴᳜≀ࡣᑡ࡞࠸ࡢ࠿ࡶࡋࢀ࡞࠸ࠋ

㸲⳦ᚑᒓᰤ㣴᳜≀࡛༟㉺ࡍࡿ⮬ᐙཷ⢊๓㡯࡛ࡣ㸪࢔ࣝࣅࣀࢆ⏝࠸࡚㸪ᙜ௦ࡢ⏕⫱࡟㛵ࡋ࡚㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡟࡝ࡢࡼ࠺
࡞㞀ቨࡀ࠶ࡿࡢ࠿ࢆ⪃࠼ࡓࠋḟ࡟ࠕ✀ࠖ࡜ࡋ࡚ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋ࡚࠸ࡿࡶࡢࢆ⏝࠸࡚㸪⦾Ṫ࡟㛵ࡋ࡚㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⋓ᚓ࡟㸪࡝ࡢࡼ࠺࡞ไ⣙ࡀ࠶ࡿࡢ࠿ࢆ⪃࠼ࡓ࠸ࠋ⳦ᚑ

K. Suetsugu - 5

植物科学最前線 5:98 （2014）ᒓᰤ㣴ᛶࡢ㐍໬ࡢ฼Ⅼ࡜ࡋ࡚ࡼࡃᣲࡆࡽࢀࡿࡶࡢࡣ㸪ගྜᡂࢆࡍࡿᚲせࡀ࡞࠸ࡓࡵ㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛⏕Ꮡྍ⬟࡟࡞ࡿ࡜࠸࠺ࡶࡢ࡛࠶ࡿࠋࡋ࠿ࡋ࡞ࡀࡽᬯ࠸⎔ቃ࡛⏕άࡍࡿࡇ࡜ࡣ㸪⳦࡜ࡢ㛵ࢃࡾ௨እࡢ㠃࡛ไ⣙ࢆཷࡅࡿࡇ࡜ࡣ࡞࠸ࡢࡔࢁ࠺࠿ࠋ౛
࠼ࡤ㸪ᬯ࠸ᯘᗋ࡛ࡢ⏕άࢆᡂࡋ㐙ࡆࡿࡓࡵ࡟ࡣ㸪⦾Ṫࡶᬯ࠸⎔ቃ࡛⾜࠺ᚲせࡀ࠶ࡿࡀ㸪ࡇ࠺ࡋࡓ⎔ቃࡣࣁࢼࣂࢳ࡞࡝ࡢゼⰼᛶ᪻⹸ࡢ࡟ࡂࢃ࠸࡜ࡣ↓⦕ࡢୡ⏺࡛࠶ࡿ 㸦Herrera
1995, Herrera 1997, Lee et al. 2001㸧ࠋࡑࡢࡓࡵ㸪⳦࡟ᐤ⏕ࡍࡿ࡜࠸࠺⏕άྐࡣ㸪ⷧᬯ࠸ᯘᗋ࡛ཷ⢊ࢆ㐩ᡂࡍࡿ࡜࠸࠺ᅔ㞴ࢆ᳜≀࡟ᙉ࠸࡚࠸ࡿࠋࡑࡇ࡛ࡇࡢࡼ࠺࡞ᬯ࠸ሙᡤ࡟⏕⫱ࡍࡿ᳜≀ࡣ㸪࡝ࡢࡼ࠺࡞᪉ἲ࡛ཷ⢊ࢆ㐩ᡂࡋ࡚࠸ࡿࡢ࠿⤂௓ࡋ࡚࠸ࡃࡇ࡜࡟ࡋࡼ࠺ࠋ ⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜ཷ⢊ᵝᘧࡢ㛫࡟ఱࡽ࠿ࡢ㛵㐃ࡀ࠶ࡿࡢ࡛ࡣ࡞࠸࠿࡜࠸࠺௬ㄝࡣ௨
๓࠿ࡽᏑᅾࡋ࡚࠾ࡾ㸪Bidartondo 㸦2005㸧 ࡣ㸪㐣ཤࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ㏦⢊ᵝᘧ࡟㛵ࡍࡿሗ࿌ࢆࡶ࡜࡟㸪
ࠕᆅୗ㒊࡛⳦᰿⳦ࢆ㦄ࡋ࡚࠸ࡿ⳦
ᚑᒓᰤ㣴᳜≀ࡣ㸪ᆅୖ㒊࡛⻤ࢆࡔࡉࡎ࡟㏦⢊⪅ࢆ㦄ࡍࡇ࡜ࡣ࡛ࡁࡎ㸪⮬Ṫࢆ᥇⏝ࡋ࡚࠸ࡿࠖ
࡜࠸࠺௬ㄝࢆᥦၐࡋࡓࠋࡋ࠿ࡋ࡞ࡀࡽ⳦ᚑᒓ
ᰤ㣴᳜≀ࡢ㏦⢊ᵝᘧ࡟㛵ࡍࡿ◊✲ࡣ㸪ᾮᾐᶆᮏࢆ⏝࠸࡚㸪ᰕ㢌࡜ⵂࡢ఩⨨㛵ಀ࡞࡝࠿ࡽཷ
⢊ᵝᘧࢆ᥎ᐃࡋࡓ◊✲ࡀ࡯࡜ࢇ࡝࡛
㸦Warming 1901, Oehler 1927㸧㸪ᐇ㝿࡟㔝እほᐹ㸪⿄᥃ࡅฎ⌮࡞࡝ࢆ㏻ࡋ࡚ヲ⣽࡟ཷ⢊ᵝᘧࢆ᳨ウࡋࡓ◊✲ࡣഹ࠿࡛࠶ࡗࡓࠋࡑࡇ࡛ᡃࠎࡣ㸪ࠕ⳦ᚑᒓᰤ㣴ᛶ࡜㏦⢊ᵝᘧࡢኚ໬࡟ࡣᐦ᥋࡞
㛵ࢃࡾࡀ࠶ࡿࠖ࡜࠸࠺௬ㄝࡢୗ㸪᪥ᮏ࡟⏘ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢཷ⢊ᵝᘧࢆヲ⣽࡟᳨ウࡋᅗ ⤖ᐇᮇࡢࢶࢳ࢔ࢣࣅⷣࡢ᫬ᮇ࡟⿄᥃ࡅࢆ⾜ࡗࡓಶయ࡜⮬↛≧ែࡢಶయ ྠ⛬ᗘࡢ⤖ᐇࡀぢࡽࢀࡿ ࡓࠋࡑࡢ⤖ᯝ㸪ࢩࣗࣥࣛࣥᒓ㸪࣒ࣚ࢘ࣛࣥᒓ㸪ࢱࢿ࢞ࢩ࣐࣒ࣚ࢘ࣛࣥᒓ㸪࢖ࣔࣛࣥᒓ㸪ࣄ࣓ࣀࣖ࢞ࣛᒓ㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘ᒓ㸪࢜ࢽࣀࣖ࢞ࣛᒓ㸪ࢶࢳ࢔ࢣࣅᒓ㸪ࢺࣛ࢟ࢳࣛ
ࣥᒓ࡞࡝ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢከࡃࡀ㸪⮬ື⮬ᐙཷ⢊ࢆ㐍໬ࡉࡏ࡚࠸ࡓࡇ࡜ࡀ᫂ࡽ࠿࡟࡞
ࡗࡓࠋ ౛࠼ࡤ㸪↓ⴥ⥳ࡢࢶࢳ࢔ࢣࣅࡣ㸪ࣂࢽࣛள⛉㸪ࣂࢽࣛ㐃࡟ᒓࡋ㸪ගྜᡂࢆࡍࡿࣂࢽࣛ
࡜㏆⦕࡛࠶ࡿ 㸦Cameron & Carmen Molina 2006㸧ࠋࣂࢽࣛᒓࡢ᳜≀ࡣ㸪ᑠᆺࡢࣁࢼࣂࢳ
࡟ࡼࡗ࡚㏦⢊ࡀ⾜ࢃࢀࡿࡶࡢࡀ࡯࡜ࢇ࡝࡛㸪᪻⹸ࡢຓࡅ࡞ࡋ࡛ཷ⢊ࡀ㉳ࡇࡿ✀㢮ࡣࡈࡃࢃࡎ࠿࡛࠶ࡿ 㸦Soto Arenas 2003㸧
ࠋࢶࢳ࢔ࢣࣅࡶⰼᙧែࡣ㸪ࣂࢽࣛࡢࡑࢀ࡜㠀ᖖ࡟ࡼࡃఝ࡚࠾ࡾ㸪ࢥࣁࢼࣂࢳ࡞࡝ࡢᑠᆺࡢࣁࢼࣂࢳ࡟ࡼࡗ࡚㏦⢊ࡉࢀࡿࡇ࡜ࡀண᝿ࡉࢀࡿࠋࡋ
࠿ࡋ࡞ࡀࡽὴᡭ࡞ⰼࢆࡋ࡚࠸ࡿ࡟ࡶ࠿࠿ࢃࡽࡎ㸪ࢶࢳ࢔ࢣࣅ࡟ࡘ࠸࡚ࡣ㸪࠿࡞ࡾ᪩࠸ࢱ
࢖࣑ࣥࢢ࡛ 㸦ሙྜ࡟ࡼࡗ࡚ࡣ㛤ⰼࡍࡿ๓࡟㸧㸪ⵂᖗ࡟᱁⣡ࡉࢀࡓⰼ⢊ሢࡀࡍࡄୗࡢᰕ㢌

K. Suetsugu - 6

植物科学最前線 5:99 （2014）࡟⁥ࡾⴠࡕ㸪⮬ື⮬ᐙཷ⢊ࡀ㉳ࡇࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ 㸦Suetsugu 2013㸧ࠋ᪩࠸ࢱ࢖࣑ࣥࢢ࡛⮬ື⮬ᐙཷ⢊ࡀ㉳ࡇࡿࡇ࡜ࢆ⪃࠼ࡿ࡜㸪ࢶࢳ࢔ࢣࣅࡣ୺࡟⮬Ṫ࡛ ✀Ꮚࢆṧࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿ 㸦ᅗ 4㸧ࠋࣂࢽࣛᒓ࡛ࡣ⮬ᐙཷ⢊ࡢ✀㢮ࡀᴟࡵ࡚ᑡ࡞࠸ࡇ࡜ࢆ⪃࠼ࡿ࡜㸪ࡇࡢ஦ࡣ㸪⳦ᚑᒓᰤ㣴ᛶ࡜⮬Ṫࡢ㐍໬࡟㛵㐃ࡀ࠶ࡿྍ⬟ᛶࢆ♧၀ࡋ࡚࠸ࡿࠋ ࡋ࠿ࡋ࡞ࡀࡽ⳦ᚑᒓᰤ㣴ᛶ࡜⮬Ṫࡢ㐍໬࡟㛵㐃ᛶࡀ࠶ࡿ࠿ࢆ᫂ࡽ࠿࡟ࡍࡿࡓࡵ࡟ࡣ㸪ヲ⣽࡞⣔⤫㛵ಀࡀ᫂ࡽ࠿࡜࡞ࡗ࡚࠸ࡿ✀⩌ࢆ⏝࠸ࡿ࡯࠺ࡀ㐺ษ࡛࠶ࡿࠋࡇࢀࡲ࡛ࡢ◊✲࠿ࡽ㸪ࢩࣗࣥࣛࣥᒓ࡛ࡣ㸪↓ⴥ⥳ᛶࡀ 1 ᅇ㐍໬ࡋࡓࡇ࡜㸪↓ⴥ⥳ᛶࡢ࣐ࣖࣛࣥ࡜ࢧ࣑࢞
ࣛࣥࡣ༢⣔⤫⩌࡜࡞ࡿࡇ࡜㸪୧✀ࡢጜጒ⩌࡟ࡣᬑ㏻ⴥࢆ⏕ࡌࡿࢼࢠࣛࣥࡀ఩⨨ࡍࡿࡇ࡜㸪ࡲࡓࡇࢀࡽࡢእ⩌ࡀࢩ࡛ࣗࣥࣛࣥ࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Yukawa et al. 2002㸧
ࠋࡲࡓᏳᐃྠ఩యศᯒ࡟ࡼࡗ࡚㸪ᬑ㏻ⴥࢆᒎ㛤ࡍࡿࢩࣗࣥࣛࣥ㸪ࢼࢠࣛࣥࡶ㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶ࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Motomura et al. 2010㸧ࠋࡼࡗ࡚ࢩࣗࣥࣛࣥᒓ
ࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜᳜≀ࡢᙧ㉁㐍໬ࢆ᳨ウࡍࡿࡢ࡟ࡍࡄࢀࡓ⣔࡛࠶ࡿ࡜࠸࠼ࡿࠋ
ࡇࡢࢩࣗࣥࣛࣥᒓ࡟࠾࠸࡚ࡶ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜ཷ⢊ᵝᘧࡢ㛫࡟ࡣ㸪┦㛵ࡀぢࡽࢀࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࢼࢠࣛࣥࡢእ⩌࡜࡞ࡿࢩࣗࣥࣛࣥ࡜࡛࢝ࣥࣛࣥࡣ㸪࣑ࢶࣂࢳࡢ࣮࣮࣡࢝࡜ࢣࣈ࢝ࣁࢼࣂࢳࡢ㞝࡟ࡑࢀࡒࢀ㏦⢊ࡉࢀ㸪⮬ື⮬ᐙཷ⢊ࡶ⾜ࢃ࡞࠸ 㸦Tsuji& Kato 2010㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࡼࡾ⳦ᚑᒓᰤ㣴ᛶࢆᙉࡵ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࢼࢠࣛࣥࡸ
ⴥࢆ⏕ࡌ࡞࠸࣐ࣖࣛࣥࡸࢧ࣑࡛࢞ࣛࣥࡣ㸪୺࡟⮬ື⮬ᐙཷ⢊ࢆ⾜ࡗ࡚࠸ࡿࠋ⯆࿡῝࠸ࡇ࡜࡟㸪୰ᅜࡢࢼࢠࣛࣥࡢಶయ⩌࡛ࡣ㸪⮬ື⮬ᐙཷ⢊ࢆ⾜ࢃ࡞࠸ಶయ⩌ࡢᏑᅾࡶ☜ㄆࡉࢀ࡚࠸ࡿ 㸦Cheng et al. 2007㸧ࠋࢩࣗࣥࣛࣥᒓ࡛ࡣ㸪↓ⴥ⥳ࡢ㐍໬࡟కࡗ࡚⭉⏕⳦࠿ࡽእ⏕⳦᰿⳦࡬࡜ࣃ࣮ࢺࢼ࣮ࡢࢩࣇࢺࡀ㉳ࡇࡗ࡚࠾ࡾ㸪ࡑࡢ㐣⛬࡛୧⪅ࢆಖᣢࡍࡿ୰㛫ẁ㝵ࢆ⤒ࡿࡇ࡜ࡀศ࠿ࡗ࡚࠸ࡿࡀ 㸦Ogura-Tsujita et al. 2012㸧
㸪㏦⢊ᵝᘧࡢ㠃࡛ࡶ㸪ⴥࢆ⏕ࡌ࡞࠸✀⩌ࡢ㐍໬࡟ඛ⾜ࡋ࡚㸪୙᏶඲࡞ᙧ࡛⮬ື⮬ᐙཷ⢊ࡀ㐍໬ࡋ࡚࠸ࡿࡢࡣ⯆࿡῝࠸ࠋ ࡇ࠺ࡋࡓ⮬Ṫࡢ㐍໬ࡣᬯ࠸ᯘᗋ࡛☜ᐇ࡟⦾Ṫࡍࡿࡢ࡟ᙺ❧ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋ⳦ᚑᒓ
ᰤ㣴᳜≀ࡀ⮬ື⮬ᐙཷ⢊ࢆ᥇⏝ࡋࡸࡍ࠸࡜࠸࠺ண

ࡣࡇࢀࡲ࡛࡟ࡶᏑᅾࡋࡓࡀ

㸦Bidartondo 2005㸧㸪⣔⤫㛵ಀࡀ᫂ࡽ࠿࡜࡞ࡗ࡚࠸ࡿ✀⩌࡛㸪㔝እほᐹ࡜⿄᥃ࡅᐇ㦂ࢆ
㏻ࡋ࡚ᐇドࡋࡓ౛࡜ࡋ࡚ᮏ◊✲ࡣ౯್ࡀ࠶ࡿࠋࡲࡓ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࡞࠿࡛ࡶ㸪⮬Ṫࢆࡲࡗࡓࡃ⾜ࢃ࡞࠸࢝ࣥࣛࣥ࡜ࢩࣗࣥࣛࣥࡣ㸪ẚ㍑ⓗ᫂ࡿ࠸⎔ቃࢆዲࡳ㸪㛤ⰼ᫬ᮇࡶᯘෙࡀ㛢㙐ࡋ࡚ࡋࡲ࠺๓ࡢ෤ࡸ᪩᫓࡛࠶ࡿⅬࡶࡁࢃࡵ࡚⯆࿡῝࠸ࠋ୍⯡ⓗ࡟ࡣ㸪᪻⹸ࡀ࠶ࡲࡾάືࡋ࡚࠸࡞࠸෤ࡸ᪩᫓࡟㛤ⰼࡍࡿࡇ࡜ࡣ୙฼࡜ࡉࢀ㸪ࡇࡢࡼ࠺࡞᫬ᮇ࡟㛤ⰼࡍࡿ᳜≀ࡣ㫽࡞࡝᪻⹸௨እࡢ㏦⢊⪅࡟౫Ꮡࡍࡿሙྜࡀከ࠸ 㸦Kunitake et al. 2004, Fang etal. 2012 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪࢝ࣥࣛࣥ࡜ࢩࣗࣥࣛࣥࡣ㸪࡯࠿ࡢ㏆⦕✀࡜ྠᵝ࡟ࣁࢼࣂࢳࢆ㏦⢊⪅࡜ࡋ࡚࠸ࡿ࡟ࡶ㛵ࢃࡽࡎ㸪෤ࡸ᪩᫓࡟㛤ⰼࡍࡿࠋࡇࢀࡽࡢ㛤ⰼ᫬ᮇࡶ㸪ࡼࡾ᫂ࡿ࠸⎔ቃ࡛ཷ⢊ࢆ㐩ᡂࡍࡿࡓࡵࡢ㐺ᛂ࡛࠶ࡿྍ⬟ᛶࡀ࠶ࡿࠋ

㸳⳦ᚑᒓᰤ㣴᳜≀࡟࠾ࡅࡿ㏦⢊⪅ࡢ㔜せᛶ๓㡯࡛㏙࡭ࡓ㏻ࡾ㸪ᬯ࠸ᯘᗋ࡟⏕⫱ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀࡟࡜ࡗ࡚㸪ࢥࣁࢼࣂࢳࡸ࣑ࢶ

K. Suetsugu - 7

植物科学最前線 5:100 （2014）ࣂࢳ࡞࡝ࡢ᫂ࡿ࠸⎔ቃࢆዲࡴ᪻⹸࡟㏦⢊ࢆ㢗ࡿࡇ࡜ࡣ㠀ᖖ࡟㞴ࡋ࠸࡜⪃࠼ࡽࢀࡿࠋࡋ࠿
ࡋ࡞ࡀࡽ㸪ᬯ࠸⎔ቃ࡟㐍ฟྍ⬟࡞᪻⹸࡟㏦⢊ࢆクࡍࡇ࡜ࡀ࡛ࡁࢀࡤ㸪ᬯ࠸ᯘᗋ࡛ࡶ௚Ṫࢆ⾜࠺ࡇ࡜ࡀྍ⬟࡛ࡣ࡞࠸ࡔࢁ࠺࠿ࠋ౛࠼ࡤ࣐ࣝࣁࢼࣂࢳࡣ㸪యㄪ⠇ࡀྍ⬟࡞ࡓࡵ㸪ᬯ࠸⎔ቃ࡛ࡶ᥇㣵ࢆ⾜࠺ഴྥࡀ࠶ࡿࠋࡋࡓࡀࡗ࡚㸪࣐ࣝࣁࢼࣂࢳࢆ࿧ࡧᐤࡏࡿࡇ࡜ࡀ࡛
ࡁࢀࡤ㸪௚Ṫ࡛⏕άྐࢆᅇࡍࡇ࡜ࡶྍ⬟࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋ஦ᐇ㸪ࢠࣥࣜࣙ࢘ࢯ࢘ࡸ
ࣔ࢖࣡ࣛࣥࢆྵࡴ࠸ࡃࡘ࠿ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪኱㔞ࡢ⻤ࢆሗ㓘࡜ࡍࡿࡇ࡜࡛㸪ࢺ࣐ࣛࣝࣁࢼࣂࢳࢆึࡵ࡜ࡍࡿ࣐ࣝࣁࢼࣂࢳࢆ㏦⢊⪅࡜ࡋ࡚฼⏝ࡋ㸪⮬ື⮬ᐙཷ⢊ࡶ⾜ࢃ࡞࠸㸦Ushimaru & Imamura 2002, ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓࢩࣙ࢘࢟ࣛࣥࡣ㸪⻤ࢆࡔࡍࡇ࡜
࡞ࡃ㸪㩭ࡸ࠿࡞ⰼࢆࡘࡅ㸪᥇㣵⤒㦂ࡢὸ࠸࣐ࣝࣁࢼࣂࢳ࡟㣵ࡀ࠶ࡿ࡜ᛮࢃࡏࡿࡇ࡜࡛㸪㏦⢊ࢆ㐩ᡂࡋ࡚࠸ࡿ 㸦ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡉࡽ࡟⳦ᚑᒓᰤ㣴᳜≀ࡢ୰࡟ࡣ㸪ࡇࢀࡲ࡛ゼⰼᛶ࡜⪃࠼ࡽࢀ࡚࠸࡞࠿ࡗࡓ᪻⹸ࢆ㏦⢊⪅࡜ࡋ࡚฼⏝ࡍࡿࡶࡢࡲ࡛Ꮡᅾࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋࡇࡢࡼ࠺࡞౛࡜ࡋ࡚㸪ᡃࠎࡣ㸪࢜ࢽࣀࣖ࢞ࣛᒓࣖࢶࢩࣟࣛࣥ⠇ࡢ≉Ṧ࡞ཷ⢊ᵝᘧࢆⓎぢࡋࡓ 㸦ᮎḟࡽ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࢩࣙ࢘ࢪࣙ࢘ࣂ࢚ࡣ㸪⭉ᩋࡀጞࡲࡗࡓᯝᐇࡸ࢟ࣀࢥࢆ᥈ࡋ࡚ᬯ࠸ᯘᗋࢆ㣕ࡧᅇࡿࡀ㸪ࣖࢶࢩࣟࣛࣥ⠇ࡢከࡃࡢ✀ࡣ㸪࢟ࣀࢥ⮯ࡸⓎ㓝⮯࡟ᨃែࡍࡿࡇ࡜࡛㸪ࢩࣙ࢘ࢪࣙ࢘ࣂ࢚࡟㏦⢊ࢆクࡋ࡚࠸ࡿࠋࡲࡓ⯆࿡῝࠸ࡇ࡜࡟ࡇࡢࣖࢶࢩࣟࣛࣥ⠇ࡢ✀⩌࡟ࡣ㸪㏆ࡃ࡟࢟ࣀࢥࡀᏑᅾࡍࡿࡇ࡜࡛㸪ཷ⢊⋡ࡀୖࡀࡿ✀㢮ࡀ࠶ࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗࡓࠋࣖࢶ
ࢩࣟࣛࣥࡣ⭉⏕⳦࡟ᐤ⏕ࡍࡿ≉ᚩ࠿ࡽ㸪ᬯࡃ‵ࡗࡓ⎔ቃ࡟⏕⫱ࡍࡿࡇ࡜ࡀከࡃ㸪஦ᐇ㸪ࣖࢶࢩࣟࣛࣥࡢ⏕࠼ࡿ࿘㎶࡟ࡣ⭉⏕⳦ࡢᏊᐇయࡀฟ⌧ࡍࡿࡇ࡜ࡶከ࠸ࠋࡘࡲࡾࣖࢶࢩࣟ
ࣛࣥࡣ㸪⮬㌟ࡢࡑࡤ࡟ከࡃ⏕࠼ࡿࠕ࢟ࣀࢥࠖ࡟㏦⢊⪅ࢆㄏᘬࡋ࡚ࡶࡽ࠺ࡇ࡜ࡀྍ⬟࡞ࡢ࡛࠶ࡿ 㸦Suetsugu & Kato 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࣖࢶࢩࣟࣛࣥࡣ㸪࿘㎶࡟࢟ࣀࢥࡀ࡞࠸⎔ቃ࡛ࡶ㸪ཷ⢊ࢆ㐩ᡂࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪࢟ࣀࢥ࡟ࡼࡿ㏦⢊⪅ࡢㄏᘬ࡟᏶඲࡟㢗ࡗ࡚࠸ࡿࢃࡅ࡛ࡣ࡞࠸ࡶࡢࡢ㸪ࣖࢶࢩࣟࣛࣥࡣ㸪ᰤ㣴ࡢࡳ࡞ࡽࡎ㏦⢊ࢧ࣮ࣅࢫࡶ㸪࢟ࣀࢥ࡟౫Ꮡࡋ࡚࠸ࡿ࡜࠸࠼ࡿࠋ
 Bidartondo 㸦2005㸧 ࡣ㸪ࠕࡍ࡛࡟ᆅୗ㒊࡛⳦᰿⳦ࢆ㦄ࡋ࡚࠸ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ᆅୖ㒊࡛⻤ࢆࡔࡉࡎ࡟㏦⢊⪅ࢆ㦄ࡍࡇ࡜ࡣ࡛ࡁ࡞࠸ࠖ࡜࠸࠺௬ㄝࢆᥦၐࡋࡓࡀ㸪ࢩࣙ࢘࢟ࣛࣥࡶࣖࢶࢩࣟࣛࣥࡶ⻤ࢆฟࡉ࡞࠸ࡓࡵ㸪ࡇࡢ௬ㄝ࡟ࡣヱᙜࡋ࡞࠸౛እࡶ࠶ࡿ࡜ゝ࠼ࡿࠋࡋ࠿ࡋ࡞ࡀࡽ㸪⻤࡞࡝ࡢሗ㓘ࢆ㏦⢊⪅࡟ᥦ౪ࡋ࡞࠸ 㸦௨ୗ↓ሗ㓘ⰼ࡜㏙࡭ࡿ㸧 ⳦ᚑᒓᰤ㣴᳜≀࡛ࡣ㸪≉࡟⮬Ṫࡀ༟㉺ࡋ࡚࠸ࡿࡢࡣ஦ᐇ࡛࠶ࡿࠋࡇࡢࡇ࡜࡟ࡣ㸪↓ሗ㓘ⰼࡣ㏦⢊⪅࡟ሗ㓘ࢆᥦ౪ࡋ࡞࠸ࡓࡵ㸪⊂❧ᰤ㣴ᛶ᳜≀ࡢሙྜ࡛ࡶ㏦⢊⪅ࡀゼࢀࡿ㢖ᗘࡀᴟࡵ࡚
ప࠸ࡇ࡜ࡀ࠾ࡑࡽࡃ㛵ಀࡋ࡚࠸ࡿ 㸦Neiland & Wilcock 1998, Tremblay et al. 2005㸧ࠋࡘࡲࡾඖࠎእ஺㓄ࡢᶵ఍ࡀᑡ࡞࠿ࡗࡓ↓ሗ㓘ⰼࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ♽ඛ✀ࡣ㸪ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋᬯ࠸⎔ቃ࡟㐍ฟࡍࡿ㐣⛬࡛㸪ࡼࡾᙉ࠸㏦⢊⪅ไ㝈ࢆࡇ࠺ࡴࡿࡇ࡜࡟࡞ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋࡑࡢࡼ࠺࡞᮲௳ୗ࡛ࡣ㸪⦾Ṫࡢಖ㞀࡜ࡋ࡚⮬ື⮬ᐙཷ⢊࡛ࡁࡿᛶ㉁ࡀ㸪ࡼࡾ᭷฼࡟ാࡃࡓࡵ㸪⻤ࢆฟࡉ࡞࠸⳦ᚑᒓᰤ㣴᳜≀࡛ࡣ㸪⮬Ṫࡀ༟㉺ࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋ

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植物科学最前線 5:101 （2014） ࡇࢀࡲ࡛⳦ᚑᒓᰤ㣴᳜≀࡛⮬Ṫࡀ༟㉺ࡍࡿせᅉ࡜ࡋ࡚ࡣ㸪ᐤ⏕⏕ά࡟క࠺㈨※ไ㝈ࡀཎᅉ࡜࠸ࢃࢀ࡚ࡁࡓ 㸦Takahashi et al. 1993, Zhang & Saunders 2000㸧
ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪㸦1㸧⮬Ṫࢆ⾜࠺⳦ᚑᒓᰤ㣴᳜≀࡛ࡶ㩭ࡸ࠿࡞ⰼࢆᣢࡘࡶࡢࡀከ࠸ࡇ࡜ 㸦Zhou et al. 2012,
Suetsugu 2013; ᅗ 5㸧 ࡸ㸪
㸦2㸧 ᬯ࠸⎔ቃ࡟⏕ᜥࡍࡿ᪻⹸ࢆ㏦⢊⪅࡟᥇⏝ࡋ࡚࠸ࡿ⳦ᚑ
ᒓᰤ㣴᳜≀࡛ࡣ㸪⮬Ṫࢆ⾜ࢃ࡞࠸ࡇ࡜ࡶከ࠸ࡇ࡜ࢆ⪃࠼ྜࢃࡏࡿ࡜ 㸦Hentrich et al. 2010,
Klooster & Culley 2009㸧
㸪㈨※ไ㝈ࡔࡅ࡛ࡣ࡞ࡃ㸪ග⎔ቃࡶ⳦ᚑᒓᰤ㣴᳜≀ࡢ⦾Ṫᵝᘧ࡟ᙳ㡪ࢆ୚࠼࡚࠸ࡿ࡜࠸࠼ࡿࠋࡇࡢࡇ࡜ࢆ⿬௜ࡅࡿドᣐ࡜ࡋ࡚ࡣ㸪࢜ࢽࣀࣖ࢞ࣛᒓࡢ࢜ࢽ
ࣀࣖ࢞ࣛࡣ㸪ᬯ࠸⎔ቃ࡛ࡣᑓࡽ࢔࣏࣑ࢡࢩࢫ࡟ࡼࡾ✀Ꮚࢆ⏕⏘ࡍࡿࡀ㸪᫂ࡿ࠸ࢠࣕࢵࣉ
࡞࡝࡟⏕⫱ࡍࡿಶయ࡛ࡣ㸪࠿࡞ࡾࡢ๭ྜ࡛ࢥࣁࢼࣂࢳ࡟ࡼࡿ௚Ṫࢆ⾜ࡗ࡚࠸ࡿࡇ࡜ࡀᣲࡆࡽࢀ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓᐟ୺࡜࡞ࡿ⳦ࡢࣁࣅࢱࢵࢺ㑅ዲᛶࡶᙳ㡪ࡍࡿࡓࡵ୍ᴫ࡟ࡣゝ࠼࡞࠸ࡀ㸪ࣁࢼࣂࢳ፹ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ᫂ࡿ࠸⎔ቃ࡟⏕⫱ࡍࡿሙྜࡀከ࠸ 㸦Suetsugu & Kato 2012, ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ⳦ᚑᒓᰤ㣴ᛶࡢ⋓ᚓ࡟ࡼࡾ㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛⏕Ꮡྍ⬟࡟࡞ࡿ࡜ࡉࢀ࡚ࡁࡓࡀ㸪ᐇ㝿ࡢ࡜ࡇࢁࡣ㸪㏦⢊ᵝᘧࡢⅬ࡛ࡶ≉ู࡞㐺ᛂࢆ㐙ࡆ࡞ࡅࢀࡤ㸪ᬯᡤ࡬ࡢ㐍ฟࡣ୙ྍ⬟࡞ࡢ࡛࠶ࡿࠋᅗ 㸬⮬ື⮬ᐙཷ⢊ྍ⬟࡞⳦ᚑᒓᰤ㣴᳜≀㸦㩭ࡸ࠿࡞ⰼࢆࡘࡅࡿࡶࡢࡀከࡃ㸪୰࡟ࡣ኱㔞ࡢⰼ⻤ࢆศἪࡍࡿࡶࡢࡶᏑᅾࡋ㸪௚Ṫࡢྍ⬟ᛶࢆ᏶඲࡟ᨺᲠࡋ࡚࠸ࡿヂ࡛ࡣ࡞࠸ࡇ࡜ࢆ᝿
㉳ࡉࡏࡿ  ୖẁᕥ࠿ࡽ㡰࡟㸪࣍ࢡࣜࢡ࣒ࣚ࢘ࣛࣥ㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘㸪ࢩࣟࢩࣕࢡࢪ
ࣙ࢘㸪࣐ࣖࣛࣥ㸪ࢶࢳ࢔ࢣࣅ㸪ࢱࢩࣟࣛࣥ

K. Suetsugu - 9

植物科学最前線 5:102 （2014）ࡲࡓ࢜ࢽࣀࣖ࢞ࣛࡢࡼ࠺࡟㸪⮬Ṫ࡜௚Ṫࢆేࡏᣢࡘ⦾Ṫᵝᘧࡣ㸪ከࡃࡢ⳦ᚑᒓᰤ㣴᳜≀࡟࡜ࡗ࡚㔜せ࡞ᙺ๭ࢆᯝࡓࡋ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ 㸦ᅗ 6㸧ࠋ⳦ᚑᒓᰤ㣴᳜≀࡜ࡑࡢ⳦᰿⳦ࡢ㛵ಀࡣ㸪ᐤ⏕⪅࡜ᐟ୺ࡢ㛵ಀ࡜࡜ࡽ࠼ࡿࡇ࡜ࡀ࡛ࡁࡿࠋࡇࢀࡲ
࡛ࡢ◊✲ࡢከࡃࡀ㸪⳦᰿ඹ⏕࡟࠾ࡅࡿඹ⏕࠿ࡽᐤ⏕࡬ࡢኚ໬࡟ࡣ≉ᐃࡢ⳦࡬ࡢ≉Ṧ໬ࡀᚲせ࡛࠶ࡿࡇ࡜ࢆ ♧ ၀ ࡋ ࡚ ࠸ ࡿ 㸦 Leake 2004,
Bidartondo 2005㸧ࠋࡇࡢඹ⏕⳦ࡢࢩ ᅗ ࢜ࢽࣀࣖ࢞ࣛ࡜ࡑࡢ㏦⢊⪅ࡢࢥࣁࢼࣂࢳ

ࣇࢺࡸ≉Ṧ໬ࡢ㉳ࡇࡿ⌮⏤࡟㛵ࡍࡿ᭷ຊ࡞௬ㄝࡀ㸪࡯࠿ࡢᐤ⏕⪅ʊᐟ୺⣔࡜ྠᵝ࡟㸪⳦ᚑᒓᰤ㣴᳜≀࡜ᐟ୺࡜ࡢ㛫࡛㌷ᣑ➇தࡀാ࠸࡚࠸ࡿ࠿ࡽ࡜࠸࠺ࡶࡢ࡛࠶ࡿ 㸦Leake
2004, Selosse & Roy 2009㸧ࠋࡇࡢࡼ࠺࡞ᩛᑐⓗ࡞┦஫స⏝ࡢሙ࡛ࡣ㸪ྠࡌ㐺ᛂᗘࢆ⥔ᣢࡍࡿࡓࡵ࡟㸪┦ᡭ࡟ᑐࡋ࡚⤯࠼ࡎ㐺ᛂࡋ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽ࡞࠸ࠋࡇࡢࡇ࡜ࡣ㸪ࣝ࢖ࢫ࣭࢟ࣕࣟࣝࡢ❺ヰࠕ㙾ࡢᅜࡢ࢔ࣜࢫࠖ࡟Ⓩሙࡍࡿ㉥ࡢዪ⋤ࡢࠕྠࡌሙᡤ࡟࡜࡝ࡲࡿࡓࡵ࡟
ࡣ㸪඲㏿ຊ࡛㉮ࡾ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽ࡞࠸ࠖ࡜࠸࠺ࢭࣜࣇ࡟࡞ࡒࡽ࠼࡚㉥ࡢዪ⋤௬ㄝ࡜࿧ࡤࢀ࡚࠸ࡿ 㸦Ladle 1992㸧ࠋࡑࡋ࡚㐺ᛂ㐍໬ࡢ㏿ᗘࢆ᪩ࡵࡿࡼ࠺࡞ᛶ㉁ࡀ᭷ᛶ⏕Ṫ࡛࠶ࡾ㸪ᩛᑐⓗ࡞┦஫స⏝ࡢᏑᅾࡇࡑࡀ㸪᭷ᛶ⏕ṪࡢᏑᅾព⩏ࢆᢸಖࡍࡿ㔜せ࡞せᅉ࡜ぢ࡞ࡉࢀ࡚࠸ࡿ 㸦Ladle 1992㸧ࠋᮏ᮶㸪⳦᰿ඹ⏕⣔࡛ࡣ㸪࠾஫࠸ࡀⰋ࠸ࣃ࣮ࢺࢼ࣮࠿࡝࠺࠿ࢆぢศࡅࡿ௙⤌ࡳࡀ࠶ࡾ㸪ࡼࡾከࡃ❅⣲ࡸࣜࣥࢆᥦ౪࡛ࡁࡿ⳦㸪ྠࡌࡃ㸪ࡼࡾከࡃⅣ⣲ࢆ౪⤥࡛ࡁࡿ᳜≀ࡀ㸪┦ᡭ࡟㑅ࡤࢀࡿ࡜ࡉࢀࡿ 㸦Kiers et al. 2011㸧ࠋࡘࡲࡾ᳜≀ࡶ⳦ࡶࣃ࣮ࢺࢼ࣮ࡢၿࡋᝏࡋࢆࡋ
ࡗ࠿ࡾ࡜㆑ูࡋ࡚࠾ࡾ㸪ཷࡅྲྀࡿ㈨※࡟ᛂࡌ࡚㏦ࡾ㎸ࡴ㣴ศࡢ㔞ࢆㄪ⠇ࡋ࡚࠸ࡿࡢ࡛࠶
ࡿࠋࣛࣥ⛉᳜≀࡜ࡑࡢ⳦᰿⳦ࡢ㛫࡛ࡣ㸪ࡇࡢࡼ࠺࡞ࠕᑂၥࠖࡢ௙⤌ࡳࡣ᫂ࡽ࠿࡟࡞ࡗ࡚
࠸࡞࠸ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࣛࣥ⛉᳜≀࡛ࡶከࡃࡢሙྜࡣ㸪ⓎⱆᙜึࡣⅣ⣲ࢆྵࡴ㣴ศࢆ⳦࡟౫Ꮡࡍࡿࡶࡢࡢ㸪⥳ⴥࢆᒎ㛤ࡋ࡚࠿ࡽⅣ⣲ࢆ⳦࡟㏉㑏ࡍࡿࡇ࡜࡛㸪┦฼ඹ⏕ࢆႠࢇ࡛࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ 㸦Cameron et al. 2006, 2008, Hynson et al. 2009㸧ࠋ࡟ࡶ࠿࠿ࢃࡽࡎ
⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ᮏ᮶㸪᳜≀ഃࡢᮏ᮶ࡢᥦ౪≀࡛࠶ࡿⅣ⣲ࢆ୚࠼࡞࠸ࡔࡅ࡛ࡣ࡞ࡃ㸪㏫࡟ዣ࠸ྲྀࡗ࡚⏕⫱ࢆྍ⬟࡟ࡋ࡚࠸ࡿࠋࡼࡗ࡚ࡑࡢ⫼ᬒ࡟ࡣ㸪ඹ⏕ࢩࢢࢼࣝࡢᨃែ࡞࡝
ࡢᕦጁ࡞ࢩࢫࢸ࣒ࡀᏑᅾࡍࡿࡇ࡜ࡀண᝿ࡉࢀࡿࠋ㉥ࡢዪ⋤௬ㄝ࡟ೌࡗ࡚⪃࠼࡚ࡳࡿ࡜㸪ᐟ୺࡜࡞ࡿ⳦㢮ࢆ㦄ࡋ⥆ࡅࡿࡓࡵ࡟ࡣ㸪⤯࠼ࡎ㐺ᛂࡋ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽࡎ㸪ᑓࡽ⮬Ṫࡢࡳࢆ⾜࠺⏕άྐࡣᴟࡵ࡚୙฼࡜⪃࠼ࡽࢀࡿࠋࡲࡓ㸪ࡇࡢࡼ࠺࡞ᐤ⏕⪅ʊᐟ୺⣔࡟㛵ࢃࡿၥ㢟Ⅼࢆ⪃࠼ࡎ࡜ࡶ㸪⮬ᐙཷ⢊ࡢࡳ࡟㢗ࡿ⏕άྐ࡟ࡣ㸪㏆஺ᙅໃࢆึࡵ࡜ࡍࡿᵝࠎ࡞୙฼┈ࡀᏑᅾࡍࡿ 㸦Lloyd 1992㸧ࠋࡇࡢࡼ࠺࡞୙฼┈ࢆゎᾘࡍࡿࡢ࡟㸪⛥࡟㉳ࡇࡿ௚Ṫࡀ

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植物科学最前線 5:103 （2014）㔜せ࡞ാࡁࢆᯝࡓࡋ࡚
࠾ࡾ㸪⮬Ṫ࡜௚Ṫࢆేࡏᣢࡘ⦾Ṫᵝᘧࡀ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕άࢫࢱ࢖ࣝ࡟࡜ࡗ࡚ࡣ㐺ࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿࠋ
ࡋ࠿ࡋ࡞ࡀࡽᴟ➃࡟ᬯᡤ࡟㐍ฟࡋࡓ⳦ᚑᒓᰤ
㣴᳜≀ࡣ㸪㛤ⰼࡍࡿࡶࡢ࡛ࡶ௚Ṫࡢᶵ఍ࢆ࡯
ࡰ᏶඲࡟ኻࡗ࡚࠾ࡾ㸪୰࡟ࡣࡘࡰࡳࡢ≧ែ࡛⮬ᐙཷ⢊ࡍࡿ᏶඲⮬Ṫᅗ ᏶඲⮬Ṫᆺࡢⰼ࡛࠶ࡿ㛢㙐ⰼࡢࡳࢆࡘࡅࡿࢶ࣑࣎ࣖࢶࢩࣟࣛࣥ ᕥ ࡜ࢱࢣࢩ࣐ࣖࢶࢩࣟࣛࣥ ྑ ᆺࡢ㛢㙐ⰼࡢࡳࢆࡘࡅࡿ✀㢮ࡶᏑᅾࡍࡿ 㸦Hsu et al. 2012, Suetsugu 2013, Suetsugu et al.
2013; ᅗ 7㸧ࠋࡑࡢࡼ࠺࡞✀㢮ࡣ㸪㐜࠿ࢀ᪩࠿ࢀ㸪⳦ࢆ㦄ࡍࡇ࡜ࡀ࡛ࡁ࡞ࡃ࡞ࡾ㸪⤯⁛ࡍࡿ㐠࿨࡟࠶ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋࡶࡋࡑ࠺࡛࠶ࡿ࡞ࡽࡤ㸪⳦࠿ࡽ㣴ศࢆ࠿ࡍࡵྲྀࡿ࡜࠸࠺
ᙼࡽࡢ⏕ࡁ᪉ࡣ㸪୍ぢ㠀ᖖ࡟㈼࠸ࡼ࠺࡟ぢ࠼ࡿࡶࡢࡢ㸪ࡑࡢ௚ࡢ㠃࡛ᩘࠎࡢ୙฼┈ࢆక࠺ࡶࡢ࡛࠶ࡾ㸪㐍໬ࡢ㐣⛬࡛⿄ᑠ㊰࡟㏞࠸㎸ࡳ㸪㛗࠸㐍໬ࡢ㐣⛬࡛ࡣᾘ࠼ࡺࡃ㐠࿨࡟࠶ࡿ࡜࡜ࡽ࠼ࡿࡇ࡜ࡀ࡛ࡁࡿ࠿ࡶࡋࢀ࡞࠸ࠋ

㸴ࡲ࡜ࡵ࡜௒ᚋࡢᒎᮃ
ᡃࠎࡢ◊✲ࢆྵࡴ㏆ᖺࡢ◊✲ࡢ㐍ᒎ࡟ࡼࡾ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ≉␗࡞ᛶ㉁ࢆ⋓ᚓ࣭⥔ᣢࡍࡿࡓࡵ࡟㸪ᐟ୺⳦࡟ᑐࡋ࡚ࡔࡅ࡛࡞ࡃ㸪௚ࡢ⏕≀࡜ࡢ┦஫స⏝࡟࠾࠸࡚ࡶᩘࠎ
ࡢᩘወ࡞㐺ᛂࢆ㐙ࡆ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋ୺࡟⮬Ṫࢆ⾜ࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚ࡁࡓࡓࡵ࠿㸪⳦ᚑᒓᰤ㣴ᛶࡢ✀⩌࡟࠾࠸࡚ࡣ㸪ࠕᕼᑡᛶࠖ㸪ࠕศᕸࠖ㸪ࠕከᵝ໬ࠖ࡞࡝ࡣ୺࡟ᐟ୺⳦࡜ࡢ┦஫స⏝ࡢᩥ⬦࡛ࡢࡳ㆟ㄽࡉࢀ㸪ࡑࡢ௚ࡢせᅉࡀ⪃៖ࡉࢀࡿࡇ࡜ࡣ
ᑡ࡞࠿ࡗࡓ 㸦Taylor et al. 2003a,b, Merckx & Bidartondo 2008, Bougoure et al. 2009, 2010,
Hazard et al. 2012㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ከࡃࡢ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࢆྵࡴࣛࣥ⛉ࡣ㸪㏦⢊ඹ⏕࡜࠸࠺㠃࡛ࡶ㠀ᖖ࡟≉Ṧ໬ࡋ࡚࠾ࡾ㸪≉ᐃࡢ㏦⢊⪅ࢆ࿧ࡧᐤࡏࡿࡓࡵ࡟㸪⊂≉࡞ⰼᙧែࢆ㐍໬ࡉࡏ࡚࠸ࡿࡶࡢࡀ࡯࡜ࢇ࡝࡛࠶ࡿ 㸦Cozzolino & Widmer 2005, Waterman &Bidartondo 2008, Waterman et al. 2011㸧
ࠋࡲࡓ㏦⢊⪅࡜ᐦ᥋࡞┦஫స⏝ࢆ⠏࠸࡚࠸ࡿࡓࡵ㸪⥳ⴥࢆᣢࡘ✀㢮࡛ࡣ㸪ࡑࡢᕼᑡᛶࡣ㸪㏦⢊⪅ࡢ≉Ṧ໬ࡢ⛬ᗘࡸ㸪㏦⢊⪅ࡢᕼᑡᛶ࡜኱ࡁ
ࡃ㛵ಀࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ㸦Neiland & Wilcock 1998, Dixon 2009, Phillips
et al. 2011㸧ࠋࡲࡓࣛࣥ⛉ࡢ⇿Ⓨⓗ࡞ከᵝ໬ࡢ⫼ᬒ࡟ࡣ㸪ཷ⢊ᵝᘧࡸ㏦⢊⪅ࡢࢩࣇࢺࡀ኱
ࡁࡃ㛵ࢃࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ 㸦Cozzolino & Widmer 2005㸧ࠋ⳦࠿ࡽ඲࡚ࡢ㣴ศࢆ␎ዣࡋ࡚࠸ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢࠕᕼᑡᛶࠖ㸪ࠕศᕸࠖ㸪ࠕከᵝ໬ࠖࡀ⳦࡜ࡢ㛵ࢃࡾ࡟ࡼࡾつ

K. Suetsugu - 11

植物科学最前線 5:104 （2014）ᐃࡉࢀ࡚࠸ࡿ࡜࠸࠺⪃࠼ࡣ㠀ᖖ࡟㨩ຊⓗ࡞ࡶࡢ࡛࠶ࡿࡀ㸪㏦⢊ᵝᘧ㸪✀Ꮚᩓᕸᵝᘧ㸪ࡑࡢ௚㠀⏕≀Ꮫⓗ࡞せᅉࢆ⪃៖࡟ධࢀ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢᐇែ࡟㏕ࡿࡇ࡜ࡀ㔜せ࡛࠶ࢁ࠺ࠋㅰ㎡ᮏ◊✲ࡣ㸪᪥ᮏᏛ⾡᣺⯆఍ࡢ≉ู◊✲ဨዡບ㈝ 㸦12J00602㸧 ࡢᨭ᥼ࢆཷࡅ࡚࠸ࡿࠋ
ࡲࡓ◊✲඲⯡㸪ཎ✏ᇳ➹࡟࠶ࡓࡗ࡚㈗㔜࡞࢔ࢻࣂ࢖ࢫࢆ㡬࠸ࡓᕝ໭⠜༤ኈ࡟ᚚ♩⏦ࡋୖࡆࡓ࠸ࠋࡲࡓ࢔ࣝࣅࣀࡢ࢟ࣥࣛࣥࡢ෗┿ࡣ㸪෗┿ᐙࡢᨾ᪂஭࿴ஓẶ࡟ࡈᥦ౪࠸ࡓࡔ࠸ࡓࠋ
ࡉࡽ࡟⤯⁛༴᝹᳜≀ࡀከ࠸㸪⳦ᚑᒓᰤ㣴᳜≀ࢆ◊✲ࡍࡿ࡟࠶ࡓࡾከࡃࡢ᳜≀ឡዲᐙࡢ᪉ࠎ࡟㸪⮬⏕ᆅሙᡤࡢ᱌ෆࡸ᝟ሗᥦ౪࡞࡝ࡢᙧ࡛࠾ୡヰ࡟࡞ࡗ࡚࠸ࡿࠋࡇࡇ࡟ࡍ࡭࡚ࡢ
ேࢆᣲࡆࡿࡇ࡜ࡣ࡛ࡁ࡞࠸ࡀ㸪ከᩘࡢ᪉ࠎࡢᨭ᥼ࢆཷࡅ㸪◊✲ࡀྍ⬟࡜࡞ࡗ࡚࠸ࡿࡇ࡜
ࢆグࡋ࡚ឤㅰࡋࡓ࠸ࠋ

䢢ᘬ⏝ᩥ⊩
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