0.01 Lellingeria apiculata GU387025 Lellingeria apiculata GU387021 Lellingeria apiculata AY362343

7079 Lellingeria affinis GU386982 Lellingeria affinis GU387041 55 Lellingeria apiculata AY460648

89 Lellingeria major GU476886 Lellingeria major GU476876

77 Lellingeria major GU386998 ネクタンドラ雲霧林自然保護区（コスタリカ）84 66 Lellingeria paramicola GU387014 66 Lellingeria major GU476873

69 Lellingeria bishopii HQ599522 64 Lellingeria major EF178627

90 Lellingeria humilis GU476897 Lellingeria tamandarei GU387027 のシダ植物DNAバーコード調査Lellingeria calolepis GU386979 95 Lellingeria pseudocapillaris GU386980 Lellingeria pseudocapillaris GU386983 1 1 Lellingeria oreophila2 GU386974 新田ジョエル 海老原淳 80 Lellingeria flagellipinnata GU387015 • • Alan R. Smith63 Lellingeria flagellipinnata GU387032 Lellingeria sp Lehnert 1232 EF178630 １ 2 100 Lellingeria bishopii GU387000 科博 植物； カリフォルニア大学バークレー校 77 Lellingeria bishopii GU476878 • 60 Lellingeria subsessilis AY460653 Lellingeria subsessilis GU386997 8765 Lellingeria subsessilis GU386996 Lellingeria subsessilis GU476885 Lellingeria pendula GU387036

65 Lellingeria jimenezii GU386986 調査地と方法 87 Lellingeria jimenezii GU386981 葉緑体rbcL Lellingeria ciliolepis GU387016 Lellingeria itatimensis GU387020 1338 bp 100 Lellingeria randallii GU387039 83 Lellingeria melanotrichia • 80% 原生雲霧林、20%二次林 99 Lellingeria randallii GU386992 78 Lellingeria suspensa 95 Lellingeria melanotrichia JNG3393 法（ ） 53 Lellingeria sp1 （図１） ML RAxML 69 Stenogrammitis limula 新記録？ Lellingeria melanotrichia GU386973 Lellingeria simacensis GU387001 100 Mycopteris taxifolia 71 3/5/2018 Map_of_Costa_Rica.svg• 158 ha、標高1200–1400 m 67 Mycopteris alsopteris Lellingeria simacensis GU476871 • 独特のrbcL配列 97 Enterosora trifurcata Lellingeria tenuicula GU386977 77 Ceradenia kalbreyeri 、 、 年にフィールド 100 Lellingeria brevistipes GU387018 を持つ • 2008 11 13 Moranopteris taenifolia Lellingeria brevistipes GU387030 調査 99 Alansmia smithii Lellingeria kaieteura GU386978 形態が 100 Cochlidium rostratum Lellingeria sp1 JNG3445 • Lellingeria 図１. ネクタンドラ 100 Cochlidium serrulatum Lellingeria isidrensis GU387042 を解読、他のシダ植物 Terpsichore atroviridis と類 の位置 • rbcL 94 Lellingeria suspensa GU386994 hombersleyi 83 Serpocaulon giganteum Lellingeria suspensa JNG3307 99 Serpocaulon loriciforme GenBankの94 rbcLと一緒に解 DNAバーコード（Ebihara et al. 100 Lellingeria suspensa GU387019 似しているが、本 Serpocaulon fraxinifolium 71 Serpocaulon ptilorhizon Lellingeria suspensa GU386976 析した系統樹の一部 Lellingeria suspensa GU387037 2010, Nia et al. 2017）と比較 100 Pecluma sp1 地域から報告が 結果と考察 57 Pecluma divaricata Lellingeria suspensa GU386969 Phlebodium pseudoaureum Lellingeria suspensa GU386970 ない Pleopeltis complanata 100 60 Lellingeria suspensa GU476824 Pleopeltis remota 63 Lellingeria suspensa GU476826

図２ 採集カーブ Microgramma lycopodioides Lellingeria dissimulans GU387043新記録 新種？ Polypodiaceae . 100 → or Microgramma percussa Lellingeria suprasculpta GU476908 Campyloneurum angustifolium 91 250 100 Lellingeria suprasculpta GU386987 予測する種数 計21科、68属、177種、6亜種 91 Campyloneurum sp1 91 100 Campyloneurum sphenodes 87 Lellingeria phlegmaria GU387008 が観察された。採集カーブ Campyloneurum brevifolium Lellingeria phlegmaria GU387034 200 100 Niphidium crassifolium Lellingeria depressa GU387017 Lellingeria hirsuta AY460649 ● （図２）はまだ頭打ちになって Oleandra decurrens 99 100 Tectaria antioquoiana 99 Lellingeria subimpressa HQ599528 100 Nephrolepis brownii Lellingeria sp Lehnert 718 EF178631 150 98 いないことから、未発見の種 85 Nephrolepis pectinata Lellingeria subimpressa GU386967 Lomariopsis maxonii Stenogrammitis myosuroides GU387006 Elaphoglossum decoratum 98 観察された種数 が残っている可能性が高い 94 Stenogrammitis myosuroides GU386999 file:///Volumes/Transcend/Documents/Conferences/2018%20JSPS/Map_of_Costa_Rica.svg 1/1 84 Elaphoglossum erinaceum

Richness 100 95 Lellingeria sp Rakotondrainibe 6882 HQ599526 Elaphoglossum crinitum と考えられる。新種か雑種の 76 Elaphoglossum lanceiforme 99 Stenogrammitis myosuroides GU387002 94 Stenogrammitis myosuroides GU386971 69 Elaphoglossum phoras 50 Elaphoglossum heterochroum 95 70 Stenogrammitis myosuroides GU386968 可能性がある物は計6 spp観 93 L. hombersleyiの採集地 99 Stenogrammitis myosuroides GU386988 67 Elaphoglossum brenesii 察された（右の系統樹に Elaphoglossum ciliatum Stenogrammitis myosuroides HQ599525 80 Elaphoglossum peltatum fo. flabellatum Lellingeria sp1 73 ネクタンドラ 0 100 100 Stenogrammitis saffordii GU476926 が付いている）。 100 Elaphoglossum doanense 82 Stenogrammitis saffordii EF178628 0 200 400 600 800 99 Elaphoglossum sp1 60 Stenogrammitis saffordii HQ599523 52 Elaphoglossum latifolium 97 100 Stenogrammitis hellwigii GU386990 Number of individuals 99 Elaphoglossum mitorrhizum Stenogrammitis hellwigii HQ599524 Elaphoglossum longicrure Mickelia oligarchica 100 Stenogrammitis wittigiana GU386989 図３ 最小種間遺伝的距離 98 Stenogrammitis wittigiana GU387029 . 61 Megalastrum sp1 新雑種？ 72 Stenogrammitis oosora HQ599527 58 97 Megalastrum longipilosum 90

68 Megalastrum apicale Stenogrammitis oosora GU386972 94 100 30% Megalastrum atrogriseum Nitta 2385 88 Stenogrammitis subcoriacea EF178629

A_Nectandra 100 Megalastrum atrogriseum Nitta 2046 Stenogrammitis subcoriacea HQ599529 ネクタンドラ 95 Megalastrum lunense Stenogrammitis limula GU476906 Stigmatopteris heterophlebia 86 20% n = 173 spp 87 Stenogrammitis limula JNG3387 100 Stigmatopteris sordida Stenogrammitis limula GU476903 Dryopteridaceae 51 Stigmatopteris contracta Stenogrammitis pumila GU387022 Dryopteris patula 10% 56 90 Stenogrammitis prionodes GU386991 Polybotrya gomezii 100 100 Stenogrammitis limula GU387023 100 Polybotrya alfredii 59 Olfersia cervina Lellingeria hartii GU386985 0% Didymochlaena truncatula Stenogrammitis limula AY460650 30% Diplazium carnosum Stenogrammitis hildebrandtii GU386975 タヒチ Diplazium urticifolium Galactodenia parrisiae KM218794

B_Tahiti 95 Diplazium striatastrum Galactodenia subscabra GU476860 ? 100 20% n = 150 spp Diplazium macrophyllum Terpsichore sp Leon 3651 AY460679 92 Diplazium obscurum Galactodenia subscabra AY460677 Diplazium palmense Melpomene flabelliformis GU387005 Athyriaceae 10% Diplazium lindbergii Melpomene aff moniliformis Sundue 1366 GU387010 Diplazium ceratolepis Melpomene moniliformis AY460654 61 100 Diplazium multigemmatum 100 Diplazium seemannii Melpomene albicans FJ011046 0% Melpomene moniliformis AY460655 30% 98 Diplazium gomezianum Diplazium myriomerum Melpomene erecta FJ011048

Frequency 日本 Melpomene melanostictaCampyloneurum GU387024 100 Blechnum appendiculatum C_Japan 51 Melpomene pilosissima GU386993 20% Blechnum occidentale Blechn- n = 684 spp 75 Lomariocycas werckleana Melpomene personataangusfolium GU476896 57 Austroblechnum lherminieri Melpomene aff xiphopteroides Labiak 4301 GU387026 75 Parablechnum proliferum Melpomene flabelliformis GU387028 10% aceae 79 100 Lomaridium ensiforme Melpomene erecta GU387013 Lomaridium fragile Melpomene personata GU387007 Salpichlaena volubilis Melpomene personata• GU387009 C. angusfolium 0% 100 Asplenium pseudoerectum Melpomene anazalea GU476898 30% 92 Asplenium pululahuae D_JapanSexDip Melpomene firma GU387035より葉の幅が 100 Asplenium cirrhatum 日本（有性生殖、二倍体のみ） 99 Asplenium rutaceum Melpomene occidentalis GU476877 71 Melpomene firma GU476864広い 100 Asplenium serra 20% 96 Aspleniaceae n = 253 spp 99 Asplenium dissectum Melpomene firma GU387003 Asplenium auritum 73 100 Melpomene occidentalis• 胞子が透明で GU476884 、 Asplenium cuspidatum 10% 100 99 Melpomene xiphopteridoides GU387040 51 Asplenium auriculatum 78 86 Melpomene xiphopteridoides形が整っていな GU387004 Asplenium holophlebium 61 Melpomene vernicosa GU476867 Hymenasplenium volubile 0% Melpomene vernicosa FJ011073い 100 Steiropteris decussata var. costaricensis 97 Melpomene sklenarii FJ011071 0%0% 3% 6% 9% 100 Steiropteris decussata var. decussata Genetic Distance 99 Steiropteris seemannii Melpomene albicans FJ011045 Thelypter- 66 Melpomene xiphopteridoides FJ011077 Goniopteris nicaraguensis 93 不稔雑種？ 100 → Amauropelta oaxacana Melpomene pseudonutans AY460657 バーコード（ ）の種間遺伝的距離が短 70 DNA rbcL 97 99 Amauropelta rudis Melpomene flabelliformis AY460656 idaceae Amauropelta villana Melpomene allosuroides GU387038 い種が多い（図３）。23/173種（13%）は種間距 Macrothelypteris torresiana Mycopteris attenuatissima GU476866 100 Pteris podophylla CampyloneurumTerpsichore sp1sp Sundue 1160 GU476879 100 離＝０（赤）、つまり、rbcLで種の識別ができな 54 Pteris speciosa Mycopteris aff semihirsuta MAS2010 GU476849 95 Pteris livida 100 Mycopteris aff semihirsuta MAS2010 GU476846 Pteris navarrensis Mycopteris attenuatissima GU476850 い。タヒチより種間距離が短いが、日本の有性 Pityrogramma calomelanos PolytaeniumMycopteris lineatum aff semihirsuta MAS2010 GU476923 Pteridaceae 100 Mycopteris leucosticta GU476921 生殖の種と似ている。 54 Vittaria bradeorum 100 Radiovittaria remota Mycopteris aff semihirsuta MAS2010 GU476857 97 Radiovittaria minima Mycopteris aff subtilis Sundue Schuettpelz 1077 GU476865 タヒチが最近（0.3–1.3 Ma）できた 100 Adiantum concinnum Mycopteris longicaulis GU476840 90 Hypolepis grandis Mycopteris attenuatissima GU476863 海洋島であるため、地域外からの 100 Hypolepis repens Mycopteris leucosticta GU476848 99 Hypolepis viscosa Terpsichore sp MAS2010f GU476890 移入が独立に起こったのに対し、 Blotiella lindeniana Mycopteris praeceps GU476839 Dennstaed- 62 Hypolepis nigrescens 59 Terpsichore sp Sundue 1167 GU476883 81 Dennstaedtia werckleana Mycopteris aff semihirsuta MAS2010 GU476899 日本やコスタリカはin situで最近種 100 Dennstaedtia auriculata 新雑種？ 67 Dennstaedtia cornuta 88 Mycopteris alsopteris AY460667 aceae Mycopteris attenuatissima GU476927 分化した種が多いと考えられる。 Saccoloma nigrescens 54 100 Cyathea mutica var. mutica 84 Mycopteris aff semihirsuta MAS2010 GU476889 100 57 Cyathea mutica var. grandis 79 Mycopteris semihirsuta GU476894

100 Cyathea nigripes Mycopteris semihirsuta AY460676 Cyathea pinnula Mycopteris aff alsopteris MAS2010 GU476915 結論 Cyathea mucilagina Mycopteris aff alsopteris MAS2010 GU476835 92 100 Cyathea delgadii Nitta 837 Mycopteris aff semihirsuta MAS2010 GU476831 Cyathea delgadii Nitta 839 75 57 Mycopteris aff• alsopteris脈の周りの葉身 MAS2010 GU476836 Cyatheales Cyathea onusta 65 Mycopteris aff semihirsuta MAS2010 GU476913 • 面積の割に種多 Sphaeropteris brunei 100 99 Cyathea bicrenata Mycopteris aff alsopterisが非常に細い MAS2010 GU476838 Alsophila polystichoides Mycopteris aff alsopteris MAS2010 GU476837 様性が高い 68 74 100 Alsophila firma 51 79 Mycopteris amphidasyon GU476922 64 木製シダの幹だ だけで種同定 Alsophila erinacea var. erinacea Mycopteris amphidasyon• GU476874 • rbcL 66 Mycopteris amphidasyon GU476872 77 Lophosoria quadripinnata Mycopteris taxifolia GU476880 Dicksonia sellowiana 73 けに特異的に生 ができないケース Didymoglossum ovale ? 94 100 Mycopteris taxifolia GU476914 100 Didymoglossum angustifrons 1cm 100 73 Mycopteris longipilosaえる GU476862 97 Didymoglossum reptans が多い 51 52 Mycopteris longipilosa GU476861 100 Didymoglossum kapplerianum Mycopteris aff taxifolia Sundue Vasco 1215 GU476888 Didymoglossum ekmanii • 新種や新雑種の 100 Mycopteris zeledoniana GU387011 100 Trichomanes crispum Trichomanes trichopodium 83 Polyphlebium Mycopteris zeledoniana GU476910 82 正体を明らかにす Vandenboschia collariata 97 Mycopteris subtilis GU386984 77 Mycopteris subtilis GU476875 58 Polyphlebium diaphanum Nitta 2022 99 るためにはより細 100 Polyphlebium diaphanum Nitta 119 capillaceumMycopteris subtilis AY460678 100 100 Polyphlebium diaphanum Nitta 2377 Mycopteris cretata GU476823 Polyphlebium sp1 かいサンプリング 100 Terpsichore sp MAS2010e HQ599521 Polyphlebium capillaceum Ascogrammitis davidsmithii GU476832 Abrodictyum rigidum Ascogrammitis davidsmithii GU476830 と核マーカーが必 Hymenophyllum elegans 80 • 葉身が普通に広い 94 Ascogrammitis davidsmithii GU476911 61 Hymenophyllum lineare 78 100 Ascogrammitis davidsmithii GU387012 要 69 Hymenophyllum fragile 97 87 Ascogrammitis davidsmithii GU476834 100 Hymenophyllum trapezoidale • 川辺の岩上に着生 Hymenophyllum consanguineum Nitta 118 Ascogrammitis davidsmithii GU476841 70 Hymenophyllum consanguineum Nitta 142 99 Ascogrammitis clavigera GU476925 • １胞子嚢に通常形の胞子Ascogrammitis clathrata GU476843 58 99 Hymenophyllum semiglabrum Hymenophyllum saenzianum 87 Ascogrammitis colombiensis GU476827 Hymenophyllum sieberi 100 95 Ascogrammitisが32個 colombiensis GU476900 Hymenophyllum hirsutum Ascogrammitis athyrioides GU476856

98 68 Hymenophyllaceae 100 Hymenophyllum polyanthos 69 Ascogrammitis pichinchae GU476928 64 Hymenophyllum myriocarpum Ascogrammitis cuencana GU476851 Hymenophyllum ectocarpon 1cm 65 Ascogrammitis dilatata GU476842 99 Sticherus retroflexus →82 雑種起源の無配生 Ascogrammitis dilatata GU387033 100 Sticherus compactus 86 Sticherus hypoleucus Ascogrammitis dilatata GU476829 Osmunda regalis Polyphlebium sp1 Ascogrammitis殖種？ loxensis GU476881 Danaea carillensis Ascogrammitis loxensis GU476852 100 99 Danaea plicata 90 Ascogrammitis loxensis GU476870 100 Danaea crispa Ascogrammitis angustipes GU476891 Ascogrammitis loxensis GU386995 Danaea moritziana 文献 80 Phlegmariurus funiformis 64 Ascogrammitis pichinchensis GU476924 100 Phlegmariurus wilsonii • Nia, J. H., J.-Y. Meyer, R. Taputuarai, and C. C. Davis. 2017. Life cycle maers: DNA barcoding reveals 99 Ascogrammitis pichinchensis GU476854 95 Phlegmariurus dichaeoides contrasng community structure between fern sporophytes and gametophytesAscogrammitis nana GU476912. Ecological Palhinhaea cernua Lycopo- 99 Ascogrammitis anfractuosa GU476853 100 Selaginella oaxacana Nitta 748 Monographs 87:278–296. 99 Ascogrammitis anfractuosa GU476845 100 Selaginella oaxacana Nitta 2371 • Ebihara, A., J. H. Nia, and M. Ito. 2010. Molecular species idenficaon with rich florisc sampling: Ascogrammitis anfractuosa GU476916 100 Selaginella porphyrospora diopsida DNA barcoding the pteridophyte flora of Japan. PLoS ONE 5:e15136. 69 Selaginella diffusa Ascogrammitis anfractuosa GU476847 99 Ascogrammitis anfractuosa GU476858

謝辞 73 Ascogrammitis anfractuosa GU476919 David and Evelyne Lennee, Freddy Casllo, Paulo Labiak100 , FAS Division of Science, Research Ascogrammitis anfractuosa GU476855 日本学術振興会（科研費Ascogrammitis anfractuosa） GU476833 Compung Group at Harvard University, 98 Ascogrammitis 15K07204 anfractuosa GU476859 Ascogrammitis nana GU387031 Ceradenia intricata KM218791 Ceradenia spixiana AY460623 Ceradenia aulaeifolia AY460619 Ceradenia aulaeifolia GU476902

99 Ceradenia intonsa GU476901 97 Ceradenia sp MAS2010 GU476893 67 Ceradenia sp MAS2010 GU476892 99 Ceradenia sp Moran 7520 GU476918

62 Ceradenia kalbreyeri GU476905 59 100 Ceradenia kalbreyeri GU476887 Ceradenia aff kalbreyeri Vasco Sundue 701 GU476825

100 Ceradenia pilipes GU476869 Ceradenia fucoides GU476907 99 98 Enterosora trifurcata GU476909 60 Enterosora trifurcata GU476917 70 Enterosora trichosora GU476920

100 Enterosora percrassa GU476882 99 Zygophlebia mathewsii GU476895 73 Zygophlebia mathewsii GU476868 Zygophlebia sectifrons GU476904

53 Notogrammitis angustifolia KM218770 86 Notogrammitis heterophylla AY460629 Notogrammitis heterophylla AY460628 Micropolypodium sikkimense JF514019

100 Alansmia lanigera AY460672 99 Alansmia lanigera GU476844

85 Alansmia cultrata GU476828 100 Leucotrichum madagascariense JN654924 95 Leucotrichum schenckii AY460651 Lellingeria pseudomitchellae AY460652

84 Serpocaulon loriciforme JNG3334 90 Serpocaulon giganteum JNG3385 100 Serpocaulon fraxinifolium JNG3327 Serpocaulon ptilorhizon JNG3390