cospeciation.pdf
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- やすはる なかじゅく
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1 oevolution Ehrlich & Raven (94) utterflies and plants: a study in coevolution oevolution Fahrenholz s Rule (9) phylogeny mirrors host phylogeny ospeciation Epichloë uchnera
2 Pocket gopher hewing lice Organism Gene rea Organism Hafner & Nadler (988) ospeciation odovergence ophylogeny ladogram rooks parsimony method (rooks, 98) Reconciliation method (Page, 994) Event-based parsimony (Ronquist, 995) Jungles (harleston, 998) Phylogram ParaFit (Legendre et al., ) ayesian method (Huelsenbeck et al., ) SH-test, U-test, likelihood-ratio test H : (Reconciliation, Event-based parsimony, Jungles, ParaFit) H : (SH-test, U-test, likelihood-ratio test)
3 rooks parsimony analysis (rooks, 98) 2つの系統群の間でどれほど共種分化が起こっているのか I II III IV I II III IV 4 E V V 2つの系統樹を重ね合わせる I II III IV 形質 I II III IV V ) 寄生者についての形質マトリクス 形質 E ) 寄主についての形質マトリクス 寄生者 ) 寄生者の系統樹の各枝に番号をふる 寄主 E E ) )のデータを寄主の系統樹上で最節約配置する 共種分化解析で想定される4つの進化的イベント 時 間 I II III IV 共種分化 cospeciation 重複 duplication ソーティング sorting 寄主転換 host switch 同一寄主上での 種分化 系統樹の不一致は必ずしも寄主転換によるものではない 絶滅 extinction 完全な寄主転換 complete switch
4 Reconciliation method Page, 994 cospeciation duplication sorting host switch uplication, Sorting Horizontal associate odivergence uplication Sorting transfer Organism gene oalescence Gene duplication, Gene loss, Gene transfer polymorphism lineage sorting parasite ospeciation Within-host Sorting, parasite switch speciation extinction Organism area Vicariance Sympatry Extinction ispersal switch Randomization Reconciliation method ), randomize ) randomize uplication, Sorting switch switch 4
5 Event-based parsimony (Ronquist, 995) phylogeny phylogeny a b c a b c phylogeny a b b c Event-based parsimony a b c ospeciation uplication Sorting c u s > d e - phylogeny c < ic < u + s Switch i > h h h h 5 h 4 a b c d e Sorting ) db,c) cost: ) ea,d) cost: ) eb or c ) or ) cost: a b c e d phylogeny a b b c a b c d e Sorting ) db,c) cost: ) ea,d) cost: ) eb or c ) or ) cost: a b b c switch a b c e d phylogeny a b c e f d phylogeny f a b c d e f ost matrix down-pass phylogeny final pass phylogeny Time 5
6 Event-based parsimony Jungles (harleston, 998) switch Jungles algorithm ladogram Hafner & Nadler (998) Sub-jungles omplete jungle (One-host-per-parasite assumption)
7 Phylogram ( ParaFit (Legendre et al., ) ParaFit (Legendre et al., ) Patristic distance Patristic distance d d d d 4 d n d d d Principal coordinate analysis imension phylogeny imension phylogeny n pairwise n- n taxa p p p p p imension imension n- dimensions 7
8 ( s s Matrix parasite (- data) s Princ. coordinates Matrix Matrix parasite (- data) Matrix ( h j ) ( p i p i p i ) = h j ( h j h j p i h j p i h j p i Matrix patristic distances Matrix H P associations Matrix patristic distances Matrix / Princ. coordinates s Matrix tree Princ. coordinates = tree Princ. coordinates Matrix Fourth-corner parameters Matrix = Matrix Fourth-corner parameters ParaFitGlobal = = trace( ) Matrix randomize ParaFitGlobal statistic ParaFit Fit ParaFit H P link trace Matrix k: trace(k) = ParaFitLink(k) = trace trace(k) ParaFitLink(k) = (trace trace(k)) / (TraceMax trace) link 8
9 ayesian method (Huelsenbeck et al., ) e = (ξ, z, γ, δ) ξ = z = γ = δ = (Hafner et al., 994) λ ξ = 9. λ =.5 ayesian method H : (SH-test, U-test, etc.) (Parametric bootstrapping, likelihood-ratio test) 9
10 (Percy et al., 4) (Rønsted et al., 5) Hafner et al. (994) OI..8 harleston M (998) Jungles: a new solution to the host/parasite phylogeny reconciliation problem. Math. iosci. 49: 9. Hafner MS, Nadler S (988) Phylogenetic trees support the coevolution of parasites and their hosts. Nature : Hafner MS, Sudman P, Villablanca FX, Spradling T, emastes JW, Nadler S (994) isparate rates of molecular evolution in cospeciating hosts and parasites. Science 5: Huelsenbeck JP, Rannala, Yang Z (997) Statistical tests of host parasite cospeciation. Evolution 5: Huelsenbeck JP, Rannala, Larget () ayesian framework for the analysis of cospeciation. Evolution 54: Legendre P, esdevises Y, azin E () statistical test for host parasite coevolution. Syst. iol. 5: 7 4. Meier-Kolthoff JP, uch F, Huson H, Gker M (7) opyat: cophylogenetic analysis tool. ioinformatics : Page RM (994) Parallel phylogenies: reconstructing the history of host parasite assemblages. ladistics : Page RM (ed.) () Tangled Trees. hicago University Press. Page RM, harleston M (998) Trees within trees: phylogeny and historical associations. Trends Ecol. Evol. : Ronquist F (995) Reconstructing the history of host parasite associations using generalized parsimony. ladistics : 7 89.
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