id sid tid token lemma pos cord-346267-l08ld2cy 1 1 key key JJ cord-346267-l08ld2cy 1 2 : : : cord-346267-l08ld2cy 1 3 cord-346267-l08ld2cy cord-346267-l08ld2cy NN cord-346267-l08ld2cy 1 4 authors author NNS cord-346267-l08ld2cy 1 5 : : : cord-346267-l08ld2cy 1 6 Wertheim Wertheim NNP cord-346267-l08ld2cy 1 7 , , , cord-346267-l08ld2cy 1 8 Joel Joel NNP cord-346267-l08ld2cy 1 9 O. O. NNP cord-346267-l08ld2cy 1 10 ; ; : cord-346267-l08ld2cy 1 11 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 1 12 Pond Pond NNP cord-346267-l08ld2cy 1 13 , , , cord-346267-l08ld2cy 1 14 Sergei Sergei NNP cord-346267-l08ld2cy 1 15 L. L. NNP cord-346267-l08ld2cy 1 16 title title NN cord-346267-l08ld2cy 1 17 : : : cord-346267-l08ld2cy 2 1 Purifying purify VBG cord-346267-l08ld2cy 2 2 Selection Selection NNP cord-346267-l08ld2cy 2 3 Can Can MD cord-346267-l08ld2cy 2 4 Obscure obscure VB cord-346267-l08ld2cy 2 5 the the DT cord-346267-l08ld2cy 2 6 Ancient Ancient NNP cord-346267-l08ld2cy 2 7 Age Age NNP cord-346267-l08ld2cy 2 8 of of IN cord-346267-l08ld2cy 2 9 Viral Viral NNP cord-346267-l08ld2cy 2 10 Lineages Lineages NNPS cord-346267-l08ld2cy 2 11 date date NN cord-346267-l08ld2cy 2 12 : : : cord-346267-l08ld2cy 2 13 2011 2011 CD cord-346267-l08ld2cy 2 14 - - HYPH cord-346267-l08ld2cy 2 15 06 06 CD cord-346267-l08ld2cy 2 16 - - HYPH cord-346267-l08ld2cy 2 17 24 24 CD cord-346267-l08ld2cy 2 18 journal journal NN cord-346267-l08ld2cy 2 19 : : : cord-346267-l08ld2cy 3 1 Molecular molecular JJ cord-346267-l08ld2cy 3 2 Biology biology NN cord-346267-l08ld2cy 3 3 and and CC cord-346267-l08ld2cy 3 4 Evolution evolution NN cord-346267-l08ld2cy 3 5 DOI doi NN cord-346267-l08ld2cy 3 6 : : : cord-346267-l08ld2cy 3 7 10.1093 10.1093 CD cord-346267-l08ld2cy 3 8 / / SYM cord-346267-l08ld2cy 3 9 molbev molbev NNP cord-346267-l08ld2cy 3 10 / / SYM cord-346267-l08ld2cy 3 11 msr170 msr170 NNP cord-346267-l08ld2cy 3 12 sha sha NNP cord-346267-l08ld2cy 3 13 : : : cord-346267-l08ld2cy 4 1 6929c0cdf46c3ce1e9c963487cc83c986a2870ef 6929c0cdf46c3ce1e9c963487cc83c986a2870ef CD cord-346267-l08ld2cy 5 1 doc_id doc_id CD cord-346267-l08ld2cy 5 2 : : : cord-346267-l08ld2cy 5 3 346267 346267 CD cord-346267-l08ld2cy 5 4 cord_uid cord_uid NNS cord-346267-l08ld2cy 5 5 : : : cord-346267-l08ld2cy 5 6 l08ld2cy l08ld2cy NNP cord-346267-l08ld2cy 6 1 Statistical statistical JJ cord-346267-l08ld2cy 6 2 methods method NNS cord-346267-l08ld2cy 6 3 for for IN cord-346267-l08ld2cy 6 4 molecular molecular JJ cord-346267-l08ld2cy 6 5 dating dating NN cord-346267-l08ld2cy 6 6 of of IN cord-346267-l08ld2cy 6 7 viral viral JJ cord-346267-l08ld2cy 6 8 origins origin NNS cord-346267-l08ld2cy 6 9 have have VBP cord-346267-l08ld2cy 6 10 been be VBN cord-346267-l08ld2cy 6 11 used use VBN cord-346267-l08ld2cy 6 12 extensively extensively RB cord-346267-l08ld2cy 6 13 to to TO cord-346267-l08ld2cy 6 14 infer infer VB cord-346267-l08ld2cy 6 15 the the DT cord-346267-l08ld2cy 6 16 time time NN cord-346267-l08ld2cy 6 17 of of IN cord-346267-l08ld2cy 6 18 most most RBS cord-346267-l08ld2cy 6 19 common common JJ cord-346267-l08ld2cy 6 20 recent recent JJ cord-346267-l08ld2cy 6 21 ancestor ancestor NN cord-346267-l08ld2cy 6 22 for for IN cord-346267-l08ld2cy 6 23 many many JJ cord-346267-l08ld2cy 6 24 rapidly rapidly RB cord-346267-l08ld2cy 6 25 evolving evolve VBG cord-346267-l08ld2cy 6 26 pathogens pathogen NNS cord-346267-l08ld2cy 6 27 . . . cord-346267-l08ld2cy 7 1 However however RB cord-346267-l08ld2cy 7 2 , , , cord-346267-l08ld2cy 7 3 there there EX cord-346267-l08ld2cy 7 4 are be VBP cord-346267-l08ld2cy 7 5 a a DT cord-346267-l08ld2cy 7 6 number number NN cord-346267-l08ld2cy 7 7 of of IN cord-346267-l08ld2cy 7 8 cases case NNS cord-346267-l08ld2cy 7 9 , , , cord-346267-l08ld2cy 7 10 in in IN cord-346267-l08ld2cy 7 11 which which WDT cord-346267-l08ld2cy 7 12 epidemiological epidemiological JJ cord-346267-l08ld2cy 7 13 , , , cord-346267-l08ld2cy 7 14 historical historical JJ cord-346267-l08ld2cy 7 15 , , , cord-346267-l08ld2cy 7 16 or or CC cord-346267-l08ld2cy 7 17 genomic genomic JJ cord-346267-l08ld2cy 7 18 evidence evidence NN cord-346267-l08ld2cy 7 19 suggests suggest VBZ cord-346267-l08ld2cy 7 20 much much RB cord-346267-l08ld2cy 7 21 older old JJR cord-346267-l08ld2cy 7 22 viral viral JJ cord-346267-l08ld2cy 7 23 origins origin NNS cord-346267-l08ld2cy 7 24 than than IN cord-346267-l08ld2cy 7 25 those those DT cord-346267-l08ld2cy 7 26 obtained obtain VBN cord-346267-l08ld2cy 7 27 via via IN cord-346267-l08ld2cy 7 28 molecular molecular JJ cord-346267-l08ld2cy 7 29 dating dating NN cord-346267-l08ld2cy 7 30 . . . cord-346267-l08ld2cy 8 1 We -PRON- PRP cord-346267-l08ld2cy 8 2 demonstrate demonstrate VBP cord-346267-l08ld2cy 8 3 how how WRB cord-346267-l08ld2cy 8 4 pervasive pervasive JJ cord-346267-l08ld2cy 8 5 purifying purifying NN cord-346267-l08ld2cy 8 6 selection selection NN cord-346267-l08ld2cy 8 7 can can MD cord-346267-l08ld2cy 8 8 mask mask VB cord-346267-l08ld2cy 8 9 the the DT cord-346267-l08ld2cy 8 10 ancient ancient JJ cord-346267-l08ld2cy 8 11 origins origin NNS cord-346267-l08ld2cy 8 12 of of IN cord-346267-l08ld2cy 8 13 recently recently RB cord-346267-l08ld2cy 8 14 sampled sample VBN cord-346267-l08ld2cy 8 15 pathogens pathogen NNS cord-346267-l08ld2cy 8 16 , , , cord-346267-l08ld2cy 8 17 in in IN cord-346267-l08ld2cy 8 18 part part NN cord-346267-l08ld2cy 8 19 due due IN cord-346267-l08ld2cy 8 20 to to IN cord-346267-l08ld2cy 8 21 the the DT cord-346267-l08ld2cy 8 22 inability inability NN cord-346267-l08ld2cy 8 23 of of IN cord-346267-l08ld2cy 8 24 nucleotide nucleotide RB cord-346267-l08ld2cy 8 25 - - HYPH cord-346267-l08ld2cy 8 26 based base VBN cord-346267-l08ld2cy 8 27 substitution substitution NN cord-346267-l08ld2cy 8 28 models model NNS cord-346267-l08ld2cy 8 29 to to TO cord-346267-l08ld2cy 8 30 properly properly RB cord-346267-l08ld2cy 8 31 account account VB cord-346267-l08ld2cy 8 32 for for IN cord-346267-l08ld2cy 8 33 complex complex JJ cord-346267-l08ld2cy 8 34 patterns pattern NNS cord-346267-l08ld2cy 8 35 of of IN cord-346267-l08ld2cy 8 36 spatial spatial JJ cord-346267-l08ld2cy 8 37 and and CC cord-346267-l08ld2cy 8 38 temporal temporal JJ cord-346267-l08ld2cy 8 39 variability variability NN cord-346267-l08ld2cy 8 40 in in IN cord-346267-l08ld2cy 8 41 selective selective JJ cord-346267-l08ld2cy 8 42 pressures pressure NNS cord-346267-l08ld2cy 8 43 . . . cord-346267-l08ld2cy 9 1 We -PRON- PRP cord-346267-l08ld2cy 9 2 use use VBP cord-346267-l08ld2cy 9 3 codon codon NN cord-346267-l08ld2cy 9 4 - - HYPH cord-346267-l08ld2cy 9 5 based base VBN cord-346267-l08ld2cy 9 6 substitution substitution NN cord-346267-l08ld2cy 9 7 models model NNS cord-346267-l08ld2cy 9 8 to to TO cord-346267-l08ld2cy 9 9 infer infer VB cord-346267-l08ld2cy 9 10 the the DT cord-346267-l08ld2cy 9 11 length length NN cord-346267-l08ld2cy 9 12 of of IN cord-346267-l08ld2cy 9 13 branches branch NNS cord-346267-l08ld2cy 9 14 in in IN cord-346267-l08ld2cy 9 15 viral viral JJ cord-346267-l08ld2cy 9 16 phylogenies phylogeny NNS cord-346267-l08ld2cy 9 17 ; ; : cord-346267-l08ld2cy 9 18 these these DT cord-346267-l08ld2cy 9 19 models model NNS cord-346267-l08ld2cy 9 20 produce produce VBP cord-346267-l08ld2cy 9 21 estimates estimate NNS cord-346267-l08ld2cy 9 22 that that WDT cord-346267-l08ld2cy 9 23 are be VBP cord-346267-l08ld2cy 9 24 often often RB cord-346267-l08ld2cy 9 25 considerably considerably RB cord-346267-l08ld2cy 9 26 longer long RBR cord-346267-l08ld2cy 9 27 than than IN cord-346267-l08ld2cy 9 28 those those DT cord-346267-l08ld2cy 9 29 obtained obtain VBN cord-346267-l08ld2cy 9 30 with with IN cord-346267-l08ld2cy 9 31 traditional traditional JJ cord-346267-l08ld2cy 9 32 nucleotide nucleotide NN cord-346267-l08ld2cy 9 33 - - HYPH cord-346267-l08ld2cy 9 34 based base VBN cord-346267-l08ld2cy 9 35 substitution substitution NN cord-346267-l08ld2cy 9 36 models model NNS cord-346267-l08ld2cy 9 37 . . . cord-346267-l08ld2cy 10 1 Correcting correct VBG cord-346267-l08ld2cy 10 2 the the DT cord-346267-l08ld2cy 10 3 apparent apparent JJ cord-346267-l08ld2cy 10 4 underestimation underestimation NN cord-346267-l08ld2cy 10 5 of of IN cord-346267-l08ld2cy 10 6 branch branch NN cord-346267-l08ld2cy 10 7 lengths length NNS cord-346267-l08ld2cy 10 8 suggests suggest VBZ cord-346267-l08ld2cy 10 9 substantially substantially RB cord-346267-l08ld2cy 10 10 older old JJR cord-346267-l08ld2cy 10 11 origins origin NNS cord-346267-l08ld2cy 10 12 for for IN cord-346267-l08ld2cy 10 13 measles measle NNS cord-346267-l08ld2cy 10 14 , , , cord-346267-l08ld2cy 10 15 Ebola Ebola NNP cord-346267-l08ld2cy 10 16 , , , cord-346267-l08ld2cy 10 17 and and CC cord-346267-l08ld2cy 10 18 avian avian JJ cord-346267-l08ld2cy 10 19 influenza influenza NN cord-346267-l08ld2cy 10 20 viruses virus NNS cord-346267-l08ld2cy 10 21 . . . cord-346267-l08ld2cy 11 1 This this DT cord-346267-l08ld2cy 11 2 work work NN cord-346267-l08ld2cy 11 3 helps help VBZ cord-346267-l08ld2cy 11 4 to to TO cord-346267-l08ld2cy 11 5 reconcile reconcile VB cord-346267-l08ld2cy 11 6 some some DT cord-346267-l08ld2cy 11 7 of of IN cord-346267-l08ld2cy 11 8 the the DT cord-346267-l08ld2cy 11 9 inconsistencies inconsistency NNS cord-346267-l08ld2cy 11 10 between between IN cord-346267-l08ld2cy 11 11 molecular molecular JJ cord-346267-l08ld2cy 11 12 dating dating NN cord-346267-l08ld2cy 11 13 and and CC cord-346267-l08ld2cy 11 14 other other JJ cord-346267-l08ld2cy 11 15 types type NNS cord-346267-l08ld2cy 11 16 of of IN cord-346267-l08ld2cy 11 17 evidence evidence NN cord-346267-l08ld2cy 11 18 concerning concern VBG cord-346267-l08ld2cy 11 19 the the DT cord-346267-l08ld2cy 11 20 age age NN cord-346267-l08ld2cy 11 21 of of IN cord-346267-l08ld2cy 11 22 viral viral JJ cord-346267-l08ld2cy 11 23 lineages lineage NNS cord-346267-l08ld2cy 11 24 . . . cord-346267-l08ld2cy 12 1 One one CD cord-346267-l08ld2cy 12 2 of of IN cord-346267-l08ld2cy 12 3 the the DT cord-346267-l08ld2cy 12 4 most most RBS cord-346267-l08ld2cy 12 5 powerful powerful JJ cord-346267-l08ld2cy 12 6 forces force NNS cord-346267-l08ld2cy 12 7 shaping shape VBG cord-346267-l08ld2cy 12 8 the the DT cord-346267-l08ld2cy 12 9 human human JJ cord-346267-l08ld2cy 12 10 genome genome NN cord-346267-l08ld2cy 12 11 is be VBZ cord-346267-l08ld2cy 12 12 adaptation adaptation NN cord-346267-l08ld2cy 12 13 to to IN cord-346267-l08ld2cy 12 14 pathogens pathogen NNS cord-346267-l08ld2cy 12 15 , , , cord-346267-l08ld2cy 12 16 and and CC cord-346267-l08ld2cy 12 17 the the DT cord-346267-l08ld2cy 12 18 pathogens pathogen NNS cord-346267-l08ld2cy 12 19 that that WDT cord-346267-l08ld2cy 12 20 appear appear VBP cord-346267-l08ld2cy 12 21 to to TO cord-346267-l08ld2cy 12 22 have have VB cord-346267-l08ld2cy 12 23 exerted exert VBN cord-346267-l08ld2cy 12 24 the the DT cord-346267-l08ld2cy 12 25 strongest strong JJS cord-346267-l08ld2cy 12 26 influence influence NN cord-346267-l08ld2cy 12 27 are be VBP cord-346267-l08ld2cy 12 28 viruses virus NNS cord-346267-l08ld2cy 13 1 ( ( -LRB- cord-346267-l08ld2cy 13 2 Worobey Worobey NNP cord-346267-l08ld2cy 13 3 et et NNP cord-346267-l08ld2cy 13 4 al al NNP cord-346267-l08ld2cy 13 5 . . . cord-346267-l08ld2cy 14 1 2007 2007 CD cord-346267-l08ld2cy 14 2 ; ; : cord-346267-l08ld2cy 14 3 Emerman Emerman NNP cord-346267-l08ld2cy 14 4 and and CC cord-346267-l08ld2cy 14 5 Malik Malik NNP cord-346267-l08ld2cy 14 6 2010 2010 CD cord-346267-l08ld2cy 14 7 ) ) -RRB- cord-346267-l08ld2cy 14 8 . . . cord-346267-l08ld2cy 15 1 Genes gene NNS cord-346267-l08ld2cy 15 2 which which WDT cord-346267-l08ld2cy 15 3 bear bear VBP cord-346267-l08ld2cy 15 4 the the DT cord-346267-l08ld2cy 15 5 mark mark NN cord-346267-l08ld2cy 15 6 of of IN cord-346267-l08ld2cy 15 7 some some DT cord-346267-l08ld2cy 15 8 of of IN cord-346267-l08ld2cy 15 9 the the DT cord-346267-l08ld2cy 15 10 most most RBS cord-346267-l08ld2cy 15 11 potent potent JJ cord-346267-l08ld2cy 15 12 selective selective JJ cord-346267-l08ld2cy 15 13 forces force NNS cord-346267-l08ld2cy 15 14 detected detect VBN cord-346267-l08ld2cy 15 15 in in IN cord-346267-l08ld2cy 15 16 the the DT cord-346267-l08ld2cy 15 17 genomes genome NNS cord-346267-l08ld2cy 15 18 of of IN cord-346267-l08ld2cy 15 19 humans human NNS cord-346267-l08ld2cy 15 20 and and CC cord-346267-l08ld2cy 15 21 our -PRON- PRP$ cord-346267-l08ld2cy 15 22 relatives relative NNS cord-346267-l08ld2cy 15 23 are be VBP cord-346267-l08ld2cy 15 24 directly directly RB cord-346267-l08ld2cy 15 25 related relate VBN cord-346267-l08ld2cy 15 26 to to IN cord-346267-l08ld2cy 15 27 combating combat VBG cord-346267-l08ld2cy 15 28 RNA RNA NNP cord-346267-l08ld2cy 15 29 viruses virus NNS cord-346267-l08ld2cy 15 30 ( ( -LRB- cord-346267-l08ld2cy 15 31 Meyerson Meyerson NNP cord-346267-l08ld2cy 15 32 and and CC cord-346267-l08ld2cy 15 33 Sawyer Sawyer NNP cord-346267-l08ld2cy 15 34 2011 2011 CD cord-346267-l08ld2cy 15 35 ) ) -RRB- cord-346267-l08ld2cy 15 36 . . . cord-346267-l08ld2cy 16 1 Moreover moreover RB cord-346267-l08ld2cy 16 2 , , , cord-346267-l08ld2cy 16 3 the the DT cord-346267-l08ld2cy 16 4 ancient ancient JJ cord-346267-l08ld2cy 16 5 history history NN cord-346267-l08ld2cy 16 6 of of IN cord-346267-l08ld2cy 16 7 this this DT cord-346267-l08ld2cy 16 8 association association NN cord-346267-l08ld2cy 16 9 with with IN cord-346267-l08ld2cy 16 10 RNA RNA NNP cord-346267-l08ld2cy 16 11 viruses virus NNS cord-346267-l08ld2cy 16 12 is be VBZ cord-346267-l08ld2cy 16 13 evidenced evidence VBN cord-346267-l08ld2cy 16 14 by by IN cord-346267-l08ld2cy 16 15 a a DT cord-346267-l08ld2cy 16 16 diverse diverse JJ cord-346267-l08ld2cy 16 17 array array NN cord-346267-l08ld2cy 16 18 of of IN cord-346267-l08ld2cy 16 19 defective defective JJ cord-346267-l08ld2cy 16 20 viral viral JJ cord-346267-l08ld2cy 16 21 remnants remnant NNS cord-346267-l08ld2cy 16 22 incorporated incorporate VBN cord-346267-l08ld2cy 16 23 into into IN cord-346267-l08ld2cy 16 24 vertebrate vertebrate JJ cord-346267-l08ld2cy 16 25 genomes genome NNS cord-346267-l08ld2cy 16 26 , , , cord-346267-l08ld2cy 16 27 including include VBG cord-346267-l08ld2cy 16 28 mounting mount VBG cord-346267-l08ld2cy 16 29 support support NN cord-346267-l08ld2cy 16 30 for for IN cord-346267-l08ld2cy 16 31 endogenization endogenization NN cord-346267-l08ld2cy 16 32 of of IN cord-346267-l08ld2cy 16 33 RNA RNA NNP cord-346267-l08ld2cy 16 34 viruses virus NNS cord-346267-l08ld2cy 16 35 other other JJ cord-346267-l08ld2cy 16 36 than than IN cord-346267-l08ld2cy 16 37 Retroviridae Retroviridae NNP cord-346267-l08ld2cy 17 1 ( ( -LRB- cord-346267-l08ld2cy 17 2 Gifford Gifford NNP cord-346267-l08ld2cy 17 3 et et NNP cord-346267-l08ld2cy 17 4 al al NNP cord-346267-l08ld2cy 17 5 . . . cord-346267-l08ld2cy 18 1 2008 2008 CD cord-346267-l08ld2cy 18 2 ; ; : cord-346267-l08ld2cy 18 3 Belyi Belyi NNP cord-346267-l08ld2cy 18 4 et et FW cord-346267-l08ld2cy 18 5 al al NNP cord-346267-l08ld2cy 18 6 . . NNP cord-346267-l08ld2cy 18 7 2010 2010 CD cord-346267-l08ld2cy 18 8 ; ; : cord-346267-l08ld2cy 18 9 Gilbert Gilbert NNP cord-346267-l08ld2cy 18 10 and and CC cord-346267-l08ld2cy 18 11 Feschotte Feschotte NNP cord-346267-l08ld2cy 18 12 2010 2010 CD cord-346267-l08ld2cy 18 13 ; ; : cord-346267-l08ld2cy 18 14 Horie Horie NNP cord-346267-l08ld2cy 18 15 et et NNP cord-346267-l08ld2cy 18 16 al al NNP cord-346267-l08ld2cy 18 17 . . . cord-346267-l08ld2cy 19 1 2010 2010 CD cord-346267-l08ld2cy 19 2 ; ; : cord-346267-l08ld2cy 19 3 Taylor Taylor NNP cord-346267-l08ld2cy 19 4 et et NNP cord-346267-l08ld2cy 19 5 al al NNP cord-346267-l08ld2cy 19 6 . . NNP cord-346267-l08ld2cy 19 7 2010 2010 CD cord-346267-l08ld2cy 19 8 ) ) -RRB- cord-346267-l08ld2cy 19 9 . . . cord-346267-l08ld2cy 20 1 However however RB cord-346267-l08ld2cy 20 2 , , , cord-346267-l08ld2cy 20 3 according accord VBG cord-346267-l08ld2cy 20 4 to to IN cord-346267-l08ld2cy 20 5 molecular molecular JJ cord-346267-l08ld2cy 20 6 dating dating NN cord-346267-l08ld2cy 20 7 analyses analysis NNS cord-346267-l08ld2cy 20 8 , , , cord-346267-l08ld2cy 20 9 many many JJ cord-346267-l08ld2cy 20 10 RNA RNA NNP cord-346267-l08ld2cy 20 11 viruses virus NNS cord-346267-l08ld2cy 20 12 have have VBP cord-346267-l08ld2cy 20 13 extraordinarily extraordinarily RB cord-346267-l08ld2cy 20 14 recent recent JJ cord-346267-l08ld2cy 20 15 origins origin NNS cord-346267-l08ld2cy 20 16 ( ( -LRB- cord-346267-l08ld2cy 20 17 Holmes Holmes NNP cord-346267-l08ld2cy 20 18 2003a 2003a CD cord-346267-l08ld2cy 20 19 ) ) -RRB- cord-346267-l08ld2cy 20 20 . . . cord-346267-l08ld2cy 21 1 At at IN cord-346267-l08ld2cy 21 2 first first JJ cord-346267-l08ld2cy 21 3 glance glance NN cord-346267-l08ld2cy 21 4 , , , cord-346267-l08ld2cy 21 5 a a DT cord-346267-l08ld2cy 21 6 recent recent JJ cord-346267-l08ld2cy 21 7 introduction introduction NN cord-346267-l08ld2cy 21 8 of of IN cord-346267-l08ld2cy 21 9 many many JJ cord-346267-l08ld2cy 21 10 RNA RNA NNP cord-346267-l08ld2cy 21 11 viruses virus NNS cord-346267-l08ld2cy 21 12 into into IN cord-346267-l08ld2cy 21 13 the the DT cord-346267-l08ld2cy 21 14 human human JJ cord-346267-l08ld2cy 21 15 population population NN cord-346267-l08ld2cy 21 16 is be VBZ cord-346267-l08ld2cy 21 17 unsurprising unsurprising JJ cord-346267-l08ld2cy 21 18 . . . cord-346267-l08ld2cy 22 1 Epidemiological epidemiological JJ cord-346267-l08ld2cy 22 2 , , , cord-346267-l08ld2cy 22 3 historical historical JJ cord-346267-l08ld2cy 22 4 , , , cord-346267-l08ld2cy 22 5 and and CC cord-346267-l08ld2cy 22 6 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 22 7 approaches approach NNS cord-346267-l08ld2cy 22 8 agree agree VBP cord-346267-l08ld2cy 22 9 that that IN cord-346267-l08ld2cy 22 10 some some DT cord-346267-l08ld2cy 22 11 of of IN cord-346267-l08ld2cy 22 12 the the DT cord-346267-l08ld2cy 22 13 most most RBS cord-346267-l08ld2cy 22 14 notable notable JJ cord-346267-l08ld2cy 22 15 RNA RNA NNP cord-346267-l08ld2cy 22 16 viruses virus NNS cord-346267-l08ld2cy 22 17 ( ( -LRB- cord-346267-l08ld2cy 22 18 e.g. e.g. RB cord-346267-l08ld2cy 22 19 , , , cord-346267-l08ld2cy 22 20 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 22 21 , , , cord-346267-l08ld2cy 22 22 Worobey Worobey NNP cord-346267-l08ld2cy 22 23 et et NNP cord-346267-l08ld2cy 22 24 al al NNP cord-346267-l08ld2cy 22 25 . . NNP cord-346267-l08ld2cy 22 26 2008 2008 CD cord-346267-l08ld2cy 22 27 ; ; , cord-346267-l08ld2cy 22 28 influenza influenza NN cord-346267-l08ld2cy 22 29 A a DT cord-346267-l08ld2cy 22 30 virus virus NN cord-346267-l08ld2cy 22 31 , , , cord-346267-l08ld2cy 22 32 Taubenberger Taubenberger NNP cord-346267-l08ld2cy 22 33 et et NNP cord-346267-l08ld2cy 22 34 al al NNP cord-346267-l08ld2cy 22 35 . . NNP cord-346267-l08ld2cy 22 36 2005 2005 CD cord-346267-l08ld2cy 22 37 ; ; : cord-346267-l08ld2cy 22 38 Ebola Ebola NNP cord-346267-l08ld2cy 22 39 virus virus NN cord-346267-l08ld2cy 22 40 [ [ -LRB- cord-346267-l08ld2cy 22 41 EBOV EBOV NNP cord-346267-l08ld2cy 22 42 ] ] -RRB- cord-346267-l08ld2cy 22 43 Zaire Zaire NNP cord-346267-l08ld2cy 22 44 , , , cord-346267-l08ld2cy 22 45 Walsh Walsh NNP cord-346267-l08ld2cy 22 46 et et NNP cord-346267-l08ld2cy 22 47 al al NNP cord-346267-l08ld2cy 22 48 . . NNP cord-346267-l08ld2cy 22 49 2005 2005 CD cord-346267-l08ld2cy 22 50 ; ; : cord-346267-l08ld2cy 22 51 and and CC cord-346267-l08ld2cy 22 52 SARS SARS NNP cord-346267-l08ld2cy 22 53 coronavirus coronavirus NN cord-346267-l08ld2cy 22 54 , , , cord-346267-l08ld2cy 22 55 Hon Hon NNP cord-346267-l08ld2cy 22 56 et et NNP cord-346267-l08ld2cy 22 57 al al NNP cord-346267-l08ld2cy 22 58 . . NNP cord-346267-l08ld2cy 22 59 2008 2008 CD cord-346267-l08ld2cy 22 60 ) ) -RRB- cord-346267-l08ld2cy 22 61 emerged emerge VBD cord-346267-l08ld2cy 22 62 as as IN cord-346267-l08ld2cy 22 63 zoonoses zoonosis NNS cord-346267-l08ld2cy 22 64 within within IN cord-346267-l08ld2cy 22 65 the the DT cord-346267-l08ld2cy 22 66 last last JJ cord-346267-l08ld2cy 22 67 century century NN cord-346267-l08ld2cy 22 68 . . . cord-346267-l08ld2cy 23 1 As as IN cord-346267-l08ld2cy 23 2 one one CD cord-346267-l08ld2cy 23 3 looks look VBZ cord-346267-l08ld2cy 23 4 further further RB cord-346267-l08ld2cy 23 5 back back RB cord-346267-l08ld2cy 23 6 in in IN cord-346267-l08ld2cy 23 7 evolutionary evolutionary JJ cord-346267-l08ld2cy 23 8 time time NN cord-346267-l08ld2cy 23 9 , , , cord-346267-l08ld2cy 23 10 however however RB cord-346267-l08ld2cy 23 11 , , , cord-346267-l08ld2cy 23 12 specific specific JJ cord-346267-l08ld2cy 23 13 inconsistencies inconsistency NNS cord-346267-l08ld2cy 23 14 arise arise VBP cord-346267-l08ld2cy 23 15 . . . cord-346267-l08ld2cy 24 1 For for IN cord-346267-l08ld2cy 24 2 example example NN cord-346267-l08ld2cy 24 3 , , , cord-346267-l08ld2cy 24 4 defective defective JJ cord-346267-l08ld2cy 24 5 remnants remnant NNS cord-346267-l08ld2cy 24 6 of of IN cord-346267-l08ld2cy 24 7 ancient ancient JJ cord-346267-l08ld2cy 24 8 integrations integration NNS cord-346267-l08ld2cy 24 9 of of IN cord-346267-l08ld2cy 24 10 Filoviridae Filoviridae NNP cord-346267-l08ld2cy 24 11 , , , cord-346267-l08ld2cy 24 12 the the DT cord-346267-l08ld2cy 24 13 viral viral JJ cord-346267-l08ld2cy 24 14 family family NN cord-346267-l08ld2cy 24 15 containing contain VBG cord-346267-l08ld2cy 24 16 the the DT cord-346267-l08ld2cy 24 17 EBOV EBOV NNP cord-346267-l08ld2cy 24 18 , , , cord-346267-l08ld2cy 24 19 are be VBP cord-346267-l08ld2cy 24 20 found find VBN cord-346267-l08ld2cy 24 21 in in IN cord-346267-l08ld2cy 24 22 mammalian mammalian JJ cord-346267-l08ld2cy 24 23 lineages lineage NNS cord-346267-l08ld2cy 24 24 that that WDT cord-346267-l08ld2cy 24 25 diverged diverge VBD cord-346267-l08ld2cy 24 26 tens ten NNS cord-346267-l08ld2cy 24 27 of of IN cord-346267-l08ld2cy 24 28 millions million NNS cord-346267-l08ld2cy 24 29 of of IN cord-346267-l08ld2cy 24 30 years year NNS cord-346267-l08ld2cy 24 31 ago ago RB cord-346267-l08ld2cy 25 1 ( ( -LRB- cord-346267-l08ld2cy 25 2 Belyi Belyi NNP cord-346267-l08ld2cy 25 3 et et FW cord-346267-l08ld2cy 25 4 al al NNP cord-346267-l08ld2cy 25 5 . . . cord-346267-l08ld2cy 26 1 2010 2010 CD cord-346267-l08ld2cy 26 2 ; ; : cord-346267-l08ld2cy 26 3 Taylor Taylor NNP cord-346267-l08ld2cy 26 4 et et NNP cord-346267-l08ld2cy 26 5 al al NNP cord-346267-l08ld2cy 26 6 . . NNP cord-346267-l08ld2cy 26 7 2010 2010 CD cord-346267-l08ld2cy 26 8 ) ) -RRB- cord-346267-l08ld2cy 26 9 , , . cord-346267-l08ld2cy 27 1 even even RB cord-346267-l08ld2cy 27 2 though though IN cord-346267-l08ld2cy 27 3 molecular molecular JJ cord-346267-l08ld2cy 27 4 dating dating NN cord-346267-l08ld2cy 27 5 suggests suggest VBZ cord-346267-l08ld2cy 27 6 a a DT cord-346267-l08ld2cy 27 7 time time NN cord-346267-l08ld2cy 27 8 of of IN cord-346267-l08ld2cy 27 9 most most RBS cord-346267-l08ld2cy 27 10 recent recent JJ cord-346267-l08ld2cy 27 11 common common JJ cord-346267-l08ld2cy 27 12 ancestor ancestor NN cord-346267-l08ld2cy 27 13 ( ( -LRB- cord-346267-l08ld2cy 27 14 tMRCA tmrca UH cord-346267-l08ld2cy 27 15 ) ) -RRB- cord-346267-l08ld2cy 27 16 for for IN cord-346267-l08ld2cy 27 17 Filoviridae Filoviridae NNP cord-346267-l08ld2cy 27 18 on on IN cord-346267-l08ld2cy 27 19 the the DT cord-346267-l08ld2cy 27 20 order order NN cord-346267-l08ld2cy 27 21 of of IN cord-346267-l08ld2cy 27 22 only only RB cord-346267-l08ld2cy 27 23 thousands thousand NNS cord-346267-l08ld2cy 27 24 of of IN cord-346267-l08ld2cy 27 25 years year NNS cord-346267-l08ld2cy 27 26 ago ago RB cord-346267-l08ld2cy 27 27 ( ( -LRB- cord-346267-l08ld2cy 27 28 Suzuki Suzuki NNP cord-346267-l08ld2cy 27 29 and and CC cord-346267-l08ld2cy 27 30 Gojobori Gojobori NNP cord-346267-l08ld2cy 27 31 1997 1997 CD cord-346267-l08ld2cy 27 32 ) ) -RRB- cord-346267-l08ld2cy 27 33 . . . cord-346267-l08ld2cy 28 1 Another another DT cord-346267-l08ld2cy 28 2 inconsistency inconsistency NN cord-346267-l08ld2cy 28 3 can can MD cord-346267-l08ld2cy 28 4 be be VB cord-346267-l08ld2cy 28 5 seen see VBN cord-346267-l08ld2cy 28 6 in in IN cord-346267-l08ld2cy 28 7 the the DT cord-346267-l08ld2cy 28 8 zoonotic zoonotic JJ cord-346267-l08ld2cy 28 9 origin origin NN cord-346267-l08ld2cy 28 10 of of IN cord-346267-l08ld2cy 28 11 measles measles NNP cord-346267-l08ld2cy 28 12 virus virus NN cord-346267-l08ld2cy 28 13 ( ( -LRB- cord-346267-l08ld2cy 28 14 MeV MeV NNP cord-346267-l08ld2cy 28 15 ) ) -RRB- cord-346267-l08ld2cy 28 16 from from IN cord-346267-l08ld2cy 28 17 rinderpest rinderpest NN cord-346267-l08ld2cy 28 18 virus virus NN cord-346267-l08ld2cy 28 19 ( ( -LRB- cord-346267-l08ld2cy 28 20 RPV RPV NNP cord-346267-l08ld2cy 28 21 ) ) -RRB- cord-346267-l08ld2cy 28 22 and and CC cord-346267-l08ld2cy 28 23 peste peste NNP cord-346267-l08ld2cy 28 24 - - HYPH cord-346267-l08ld2cy 28 25 des des NNP cord-346267-l08ld2cy 28 26 - - HYPH cord-346267-l08ld2cy 28 27 petits petit NNS cord-346267-l08ld2cy 28 28 ruminants ruminant NNS cord-346267-l08ld2cy 28 29 virus virus NN cord-346267-l08ld2cy 28 30 ( ( -LRB- cord-346267-l08ld2cy 28 31 PPRV PPRV NNP cord-346267-l08ld2cy 28 32 ) ) -RRB- cord-346267-l08ld2cy 28 33 , , . cord-346267-l08ld2cy 29 1 two two CD cord-346267-l08ld2cy 29 2 viruses virus NNS cord-346267-l08ld2cy 29 3 capable capable JJ cord-346267-l08ld2cy 29 4 of of IN cord-346267-l08ld2cy 29 5 infecting infect VBG cord-346267-l08ld2cy 29 6 large large JJ cord-346267-l08ld2cy 29 7 and and CC cord-346267-l08ld2cy 29 8 small small JJ cord-346267-l08ld2cy 29 9 ruminants ruminant NNS cord-346267-l08ld2cy 29 10 , , , cord-346267-l08ld2cy 29 11 respectively respectively RB cord-346267-l08ld2cy 29 12 . . . cord-346267-l08ld2cy 30 1 MeV MeV NNP cord-346267-l08ld2cy 30 2 required require VBD cord-346267-l08ld2cy 30 3 at at RB cord-346267-l08ld2cy 30 4 least least RBS cord-346267-l08ld2cy 30 5 two two CD cord-346267-l08ld2cy 30 6 conditions condition NNS cord-346267-l08ld2cy 30 7 to to TO cord-346267-l08ld2cy 30 8 emerge emerge VB cord-346267-l08ld2cy 30 9 : : : cord-346267-l08ld2cy 30 10 1 1 LS cord-346267-l08ld2cy 30 11 ) ) -RRB- cord-346267-l08ld2cy 30 12 Humans human NNS cord-346267-l08ld2cy 30 13 had have VBD cord-346267-l08ld2cy 30 14 to to TO cord-346267-l08ld2cy 30 15 live live VB cord-346267-l08ld2cy 30 16 in in IN cord-346267-l08ld2cy 30 17 close close JJ cord-346267-l08ld2cy 30 18 proximity proximity NN cord-346267-l08ld2cy 30 19 to to IN cord-346267-l08ld2cy 30 20 RPV RPV NNP cord-346267-l08ld2cy 30 21 , , , cord-346267-l08ld2cy 30 22 which which WDT cord-346267-l08ld2cy 30 23 became become VBD cord-346267-l08ld2cy 30 24 commonplace commonplace JJ cord-346267-l08ld2cy 30 25 only only RB cord-346267-l08ld2cy 30 26 after after IN cord-346267-l08ld2cy 30 27 the the DT cord-346267-l08ld2cy 30 28 domestication domestication NN cord-346267-l08ld2cy 30 29 of of IN cord-346267-l08ld2cy 30 30 cattle cattle NNS cord-346267-l08ld2cy 30 31 over over IN cord-346267-l08ld2cy 30 32 the the DT cord-346267-l08ld2cy 30 33 last last JJ cord-346267-l08ld2cy 30 34 10,000 10,000 CD cord-346267-l08ld2cy 30 35 years year NNS cord-346267-l08ld2cy 30 36 ( ( -LRB- cord-346267-l08ld2cy 30 37 Perkins Perkins NNP cord-346267-l08ld2cy 30 38 1969 1969 CD cord-346267-l08ld2cy 30 39 ; ; : cord-346267-l08ld2cy 30 40 Loftus Loftus NNP cord-346267-l08ld2cy 30 41 et et NNP cord-346267-l08ld2cy 30 42 al al NNP cord-346267-l08ld2cy 30 43 . . . cord-346267-l08ld2cy 30 44 1994 1994 CD cord-346267-l08ld2cy 30 45 ; ; : cord-346267-l08ld2cy 30 46 Beja Beja NNP cord-346267-l08ld2cy 30 47 - - HYPH cord-346267-l08ld2cy 30 48 Pereira Pereira NNP cord-346267-l08ld2cy 30 49 et et NNP cord-346267-l08ld2cy 30 50 al al NNP cord-346267-l08ld2cy 30 51 . . . cord-346267-l08ld2cy 30 52 2006 2006 CD cord-346267-l08ld2cy 30 53 ) ) -RRB- cord-346267-l08ld2cy 30 54 and and CC cord-346267-l08ld2cy 31 1 2 2 LS cord-346267-l08ld2cy 31 2 ) ) -RRB- cord-346267-l08ld2cy 31 3 humans human NNS cord-346267-l08ld2cy 31 4 needed need VBD cord-346267-l08ld2cy 31 5 societies society NNS cord-346267-l08ld2cy 31 6 with with IN cord-346267-l08ld2cy 31 7 a a DT cord-346267-l08ld2cy 31 8 population population NN cord-346267-l08ld2cy 31 9 size size NN cord-346267-l08ld2cy 31 10 above above IN cord-346267-l08ld2cy 31 11 250,000 250,000 CD cord-346267-l08ld2cy 31 12 - - SYM cord-346267-l08ld2cy 31 13 500,000 500,000 CD cord-346267-l08ld2cy 31 14 to to TO cord-346267-l08ld2cy 31 15 sustain sustain VB cord-346267-l08ld2cy 31 16 the the DT cord-346267-l08ld2cy 31 17 epidemic epidemic NN cord-346267-l08ld2cy 31 18 , , , cord-346267-l08ld2cy 31 19 which which WDT cord-346267-l08ld2cy 31 20 did do VBD cord-346267-l08ld2cy 31 21 not not RB cord-346267-l08ld2cy 31 22 exist exist VB cord-346267-l08ld2cy 31 23 until until IN cord-346267-l08ld2cy 31 24 about about RB cord-346267-l08ld2cy 31 25 5,000 5,000 CD cord-346267-l08ld2cy 31 26 years year NNS cord-346267-l08ld2cy 31 27 ago ago RB cord-346267-l08ld2cy 31 28 ( ( -LRB- cord-346267-l08ld2cy 32 1 Black black JJ cord-346267-l08ld2cy 32 2 1966 1966 CD cord-346267-l08ld2cy 32 3 ) ) -RRB- cord-346267-l08ld2cy 32 4 . . . cord-346267-l08ld2cy 33 1 Furthermore furthermore RB cord-346267-l08ld2cy 33 2 , , , cord-346267-l08ld2cy 33 3 the the DT cord-346267-l08ld2cy 33 4 first first JJ cord-346267-l08ld2cy 33 5 unambiguous unambiguous JJ cord-346267-l08ld2cy 33 6 historical historical JJ cord-346267-l08ld2cy 33 7 account account NN cord-346267-l08ld2cy 33 8 of of IN cord-346267-l08ld2cy 33 9 measles measle NNS cord-346267-l08ld2cy 33 10 dates date VBZ cord-346267-l08ld2cy 33 11 back back RB cord-346267-l08ld2cy 33 12 to to IN cord-346267-l08ld2cy 33 13 the the DT cord-346267-l08ld2cy 33 14 ninth ninth JJ cord-346267-l08ld2cy 33 15 century century NN cord-346267-l08ld2cy 33 16 ( ( -LRB- cord-346267-l08ld2cy 33 17 Rāzī Rāzī NNP cord-346267-l08ld2cy 33 18 1848 1848 CD cord-346267-l08ld2cy 33 19 ) ) -RRB- cord-346267-l08ld2cy 33 20 . . . cord-346267-l08ld2cy 34 1 Therefore therefore RB cord-346267-l08ld2cy 34 2 , , , cord-346267-l08ld2cy 34 3 historical historical JJ cord-346267-l08ld2cy 34 4 and and CC cord-346267-l08ld2cy 34 5 epidemiological epidemiological JJ cord-346267-l08ld2cy 34 6 considerations consideration NNS cord-346267-l08ld2cy 34 7 , , , cord-346267-l08ld2cy 34 8 which which WDT cord-346267-l08ld2cy 34 9 place place VBP cord-346267-l08ld2cy 34 10 the the DT cord-346267-l08ld2cy 34 11 tMRCA tmrca NN cord-346267-l08ld2cy 34 12 of of IN cord-346267-l08ld2cy 34 13 MeV MeV NNP cord-346267-l08ld2cy 34 14 and and CC cord-346267-l08ld2cy 34 15 RPV RPV NNP cord-346267-l08ld2cy 34 16 at at IN cord-346267-l08ld2cy 34 17 thousands thousand NNS cord-346267-l08ld2cy 34 18 of of IN cord-346267-l08ld2cy 34 19 years year NNS cord-346267-l08ld2cy 34 20 ago ago RB cord-346267-l08ld2cy 34 21 , , , cord-346267-l08ld2cy 34 22 are be VBP cord-346267-l08ld2cy 34 23 at at IN cord-346267-l08ld2cy 34 24 odds odd NNS cord-346267-l08ld2cy 34 25 with with IN cord-346267-l08ld2cy 34 26 molecular molecular JJ cord-346267-l08ld2cy 34 27 dating dating NN cord-346267-l08ld2cy 34 28 analysis analysis NN cord-346267-l08ld2cy 34 29 , , , cord-346267-l08ld2cy 34 30 which which WDT cord-346267-l08ld2cy 34 31 infers infer VBZ cord-346267-l08ld2cy 34 32 the the DT cord-346267-l08ld2cy 34 33 tMRCA tmrca NN cord-346267-l08ld2cy 34 34 to to TO cord-346267-l08ld2cy 34 35 be be VB cord-346267-l08ld2cy 34 36 only only RB cord-346267-l08ld2cy 34 37 hundreds hundred NNS cord-346267-l08ld2cy 34 38 of of IN cord-346267-l08ld2cy 34 39 years year NNS cord-346267-l08ld2cy 34 40 ago ago RB cord-346267-l08ld2cy 35 1 ( ( -LRB- cord-346267-l08ld2cy 35 2 Furuse Furuse NNP cord-346267-l08ld2cy 35 3 et et NNP cord-346267-l08ld2cy 35 4 al al NNP cord-346267-l08ld2cy 35 5 . . NNP cord-346267-l08ld2cy 35 6 2010 2010 CD cord-346267-l08ld2cy 35 7 ) ) -RRB- cord-346267-l08ld2cy 35 8 . . . cord-346267-l08ld2cy 36 1 The the DT cord-346267-l08ld2cy 36 2 same same JJ cord-346267-l08ld2cy 36 3 phylogenetic phylogenetic NN cord-346267-l08ld2cy 36 4 dating dating NN cord-346267-l08ld2cy 36 5 methods method NNS cord-346267-l08ld2cy 36 6 that that WDT cord-346267-l08ld2cy 36 7 provide provide VBP cord-346267-l08ld2cy 36 8 convincing convincing JJ cord-346267-l08ld2cy 36 9 and and CC cord-346267-l08ld2cy 36 10 reasonable reasonable JJ cord-346267-l08ld2cy 36 11 tMRCA tmrca NN cord-346267-l08ld2cy 36 12 estimates estimate NNS cord-346267-l08ld2cy 36 13 for for IN cord-346267-l08ld2cy 36 14 recent recent JJ cord-346267-l08ld2cy 36 15 zoonotic zoonotic JJ cord-346267-l08ld2cy 36 16 transfers transfer NNS cord-346267-l08ld2cy 36 17 seem seem VBP cord-346267-l08ld2cy 36 18 to to TO cord-346267-l08ld2cy 36 19 be be VB cord-346267-l08ld2cy 36 20 dramatically dramatically RB cord-346267-l08ld2cy 36 21 underestimating underestimate VBG cord-346267-l08ld2cy 36 22 the the DT cord-346267-l08ld2cy 36 23 age age NN cord-346267-l08ld2cy 36 24 of of IN cord-346267-l08ld2cy 36 25 older old JJR cord-346267-l08ld2cy 36 26 viral viral JJ cord-346267-l08ld2cy 36 27 divergence divergence NN cord-346267-l08ld2cy 36 28 events event NNS cord-346267-l08ld2cy 36 29 . . . cord-346267-l08ld2cy 37 1 Therefore therefore RB cord-346267-l08ld2cy 37 2 , , , cord-346267-l08ld2cy 37 3 over over IN cord-346267-l08ld2cy 37 4 longer long JJR cord-346267-l08ld2cy 37 5 periods period NNS cord-346267-l08ld2cy 37 6 of of IN cord-346267-l08ld2cy 37 7 time time NN cord-346267-l08ld2cy 37 8 , , , cord-346267-l08ld2cy 37 9 the the DT cord-346267-l08ld2cy 37 10 extent extent NN cord-346267-l08ld2cy 37 11 of of IN cord-346267-l08ld2cy 37 12 evolutionary evolutionary JJ cord-346267-l08ld2cy 37 13 change change NN cord-346267-l08ld2cy 37 14 has have VBZ cord-346267-l08ld2cy 37 15 been be VBN cord-346267-l08ld2cy 37 16 lost lose VBN cord-346267-l08ld2cy 37 17 . . . cord-346267-l08ld2cy 38 1 In in IN cord-346267-l08ld2cy 38 2 many many JJ cord-346267-l08ld2cy 38 3 viral viral JJ cord-346267-l08ld2cy 38 4 genes gene NNS cord-346267-l08ld2cy 38 5 , , , cord-346267-l08ld2cy 38 6 there there EX cord-346267-l08ld2cy 38 7 is be VBZ cord-346267-l08ld2cy 38 8 remarkable remarkable JJ cord-346267-l08ld2cy 38 9 sequence sequence NN cord-346267-l08ld2cy 38 10 conservation conservation NN cord-346267-l08ld2cy 38 11 , , , cord-346267-l08ld2cy 38 12 indicating indicate VBG cord-346267-l08ld2cy 38 13 that that IN cord-346267-l08ld2cy 38 14 purifying purify VBG cord-346267-l08ld2cy 38 15 selection selection NN cord-346267-l08ld2cy 38 16 is be VBZ cord-346267-l08ld2cy 38 17 a a DT cord-346267-l08ld2cy 38 18 dominant dominant JJ cord-346267-l08ld2cy 38 19 evolutionary evolutionary JJ cord-346267-l08ld2cy 38 20 force force NN cord-346267-l08ld2cy 38 21 , , , cord-346267-l08ld2cy 38 22 acting act VBG cord-346267-l08ld2cy 38 23 to to TO cord-346267-l08ld2cy 38 24 maintain maintain VB cord-346267-l08ld2cy 38 25 evidence evidence NN cord-346267-l08ld2cy 38 26 of of IN cord-346267-l08ld2cy 38 27 homology homology NN cord-346267-l08ld2cy 38 28 by by IN cord-346267-l08ld2cy 38 29 preserving preserve VBG cord-346267-l08ld2cy 38 30 amino amino NN cord-346267-l08ld2cy 38 31 acid acid NN cord-346267-l08ld2cy 38 32 residues residue NNS cord-346267-l08ld2cy 38 33 , , , cord-346267-l08ld2cy 38 34 while while IN cord-346267-l08ld2cy 38 35 allowing allow VBG cord-346267-l08ld2cy 38 36 nucleotide nucleotide JJ cord-346267-l08ld2cy 38 37 sequences sequence NNS cord-346267-l08ld2cy 38 38 to to TO cord-346267-l08ld2cy 38 39 continue continue VB cord-346267-l08ld2cy 38 40 evolving evolve VBG cord-346267-l08ld2cy 38 41 MBE MBE NNP cord-346267-l08ld2cy 38 42 ( ( -LRB- cord-346267-l08ld2cy 38 43 Holmes Holmes NNP cord-346267-l08ld2cy 38 44 2003b 2003b NNS cord-346267-l08ld2cy 38 45 ; ; : cord-346267-l08ld2cy 38 46 Edwards Edwards NNP cord-346267-l08ld2cy 38 47 et et NNP cord-346267-l08ld2cy 38 48 al al NNP cord-346267-l08ld2cy 38 49 . . . cord-346267-l08ld2cy 39 1 2006 2006 CD cord-346267-l08ld2cy 39 2 ; ; : cord-346267-l08ld2cy 39 3 Pybus Pybus NNP cord-346267-l08ld2cy 39 4 et et NNP cord-346267-l08ld2cy 39 5 al al NNP cord-346267-l08ld2cy 39 6 . . NNP cord-346267-l08ld2cy 39 7 2007 2007 CD cord-346267-l08ld2cy 39 8 ) ) -RRB- cord-346267-l08ld2cy 39 9 . . . cord-346267-l08ld2cy 40 1 Although although IN cord-346267-l08ld2cy 40 2 the the DT cord-346267-l08ld2cy 40 3 effect effect NN cord-346267-l08ld2cy 40 4 of of IN cord-346267-l08ld2cy 40 5 purifying purify VBG cord-346267-l08ld2cy 40 6 selection selection NN cord-346267-l08ld2cy 40 7 on on IN cord-346267-l08ld2cy 40 8 mutations mutation NNS cord-346267-l08ld2cy 40 9 in in IN cord-346267-l08ld2cy 40 10 RNA RNA NNP cord-346267-l08ld2cy 40 11 viruses virus NNS cord-346267-l08ld2cy 40 12 is be VBZ cord-346267-l08ld2cy 40 13 becoming become VBG cord-346267-l08ld2cy 40 14 better well RBR cord-346267-l08ld2cy 40 15 understood understand VBN cord-346267-l08ld2cy 40 16 ( ( -LRB- cord-346267-l08ld2cy 40 17 Belshaw Belshaw NNP cord-346267-l08ld2cy 40 18 et et NNP cord-346267-l08ld2cy 40 19 al al NNP cord-346267-l08ld2cy 40 20 . . NNP cord-346267-l08ld2cy 40 21 2008 2008 CD cord-346267-l08ld2cy 40 22 ) ) -RRB- cord-346267-l08ld2cy 40 23 , , , cord-346267-l08ld2cy 40 24 the the DT cord-346267-l08ld2cy 40 25 importance importance NN cord-346267-l08ld2cy 40 26 of of IN cord-346267-l08ld2cy 40 27 spatial spatial JJ cord-346267-l08ld2cy 40 28 and and CC cord-346267-l08ld2cy 40 29 temporal temporal JJ cord-346267-l08ld2cy 40 30 variation variation NN cord-346267-l08ld2cy 40 31 in in IN cord-346267-l08ld2cy 40 32 selection selection NN cord-346267-l08ld2cy 40 33 pressures pressure NNS cord-346267-l08ld2cy 40 34 has have VBZ cord-346267-l08ld2cy 40 35 not not RB cord-346267-l08ld2cy 40 36 been be VBN cord-346267-l08ld2cy 40 37 explicitly explicitly RB cord-346267-l08ld2cy 40 38 explored explore VBN cord-346267-l08ld2cy 40 39 in in IN cord-346267-l08ld2cy 40 40 the the DT cord-346267-l08ld2cy 40 41 context context NN cord-346267-l08ld2cy 40 42 of of IN cord-346267-l08ld2cy 40 43 tMRCA tmrca NN cord-346267-l08ld2cy 40 44 estimation estimation NN cord-346267-l08ld2cy 40 45 . . . cord-346267-l08ld2cy 41 1 It -PRON- PRP cord-346267-l08ld2cy 41 2 has have VBZ cord-346267-l08ld2cy 41 3 been be VBN cord-346267-l08ld2cy 41 4 well well RB cord-346267-l08ld2cy 41 5 established establish VBN cord-346267-l08ld2cy 41 6 that that IN cord-346267-l08ld2cy 41 7 failing fail VBG cord-346267-l08ld2cy 41 8 to to TO cord-346267-l08ld2cy 41 9 account account VB cord-346267-l08ld2cy 41 10 for for IN cord-346267-l08ld2cy 41 11 site site NN cord-346267-l08ld2cy 41 12 - - HYPH cord-346267-l08ld2cy 41 13 to to IN cord-346267-l08ld2cy 41 14 - - HYPH cord-346267-l08ld2cy 41 15 site site NN cord-346267-l08ld2cy 41 16 rate rate NN cord-346267-l08ld2cy 41 17 variation variation NN cord-346267-l08ld2cy 41 18 can can MD cord-346267-l08ld2cy 41 19 lead lead VB cord-346267-l08ld2cy 41 20 to to IN cord-346267-l08ld2cy 41 21 an an DT cord-346267-l08ld2cy 41 22 underestimation underestimation NN cord-346267-l08ld2cy 41 23 of of IN cord-346267-l08ld2cy 41 24 branch branch NN cord-346267-l08ld2cy 41 25 lengths length NNS cord-346267-l08ld2cy 41 26 due due JJ cord-346267-l08ld2cy 41 27 to to IN cord-346267-l08ld2cy 41 28 repeated repeat VBN cord-346267-l08ld2cy 41 29 substitutions substitution NNS cord-346267-l08ld2cy 41 30 at at IN cord-346267-l08ld2cy 41 31 rapidly rapidly RB cord-346267-l08ld2cy 41 32 evolving evolve VBG cord-346267-l08ld2cy 41 33 sites site NNS cord-346267-l08ld2cy 41 34 ( ( -LRB- cord-346267-l08ld2cy 41 35 Brown Brown NNP cord-346267-l08ld2cy 41 36 et et NNP cord-346267-l08ld2cy 41 37 al al NNP cord-346267-l08ld2cy 41 38 . . NNP cord-346267-l08ld2cy 41 39 1982 1982 CD cord-346267-l08ld2cy 41 40 ; ; : cord-346267-l08ld2cy 41 41 Sullivan Sullivan NNP cord-346267-l08ld2cy 41 42 and and CC cord-346267-l08ld2cy 41 43 Joyce Joyce NNP cord-346267-l08ld2cy 41 44 2005 2005 CD cord-346267-l08ld2cy 41 45 ) ) -RRB- cord-346267-l08ld2cy 41 46 , , , cord-346267-l08ld2cy 41 47 and and CC cord-346267-l08ld2cy 41 48 the the DT cord-346267-l08ld2cy 41 49 use use NN cord-346267-l08ld2cy 41 50 of of IN cord-346267-l08ld2cy 41 51 models model NNS cord-346267-l08ld2cy 41 52 , , , cord-346267-l08ld2cy 41 53 which which WDT cord-346267-l08ld2cy 41 54 permit permit VBP cord-346267-l08ld2cy 41 55 such such JJ cord-346267-l08ld2cy 41 56 variation variation NN cord-346267-l08ld2cy 41 57 ( ( -LRB- cord-346267-l08ld2cy 41 58 e.g. e.g. RB cord-346267-l08ld2cy 41 59 , , , cord-346267-l08ld2cy 41 60 Yang Yang NNP cord-346267-l08ld2cy 41 61 1993 1993 CD cord-346267-l08ld2cy 41 62 ) ) -RRB- cord-346267-l08ld2cy 41 63 , , , cord-346267-l08ld2cy 41 64 have have VBP cord-346267-l08ld2cy 41 65 become become VBN cord-346267-l08ld2cy 41 66 standard standard JJ cord-346267-l08ld2cy 41 67 . . . cord-346267-l08ld2cy 42 1 More more RBR cord-346267-l08ld2cy 42 2 recently recently RB cord-346267-l08ld2cy 42 3 , , , cord-346267-l08ld2cy 42 4 Suchard Suchard NNP cord-346267-l08ld2cy 42 5 and and CC cord-346267-l08ld2cy 42 6 Rambaut Rambaut NNP cord-346267-l08ld2cy 42 7 ( ( -LRB- cord-346267-l08ld2cy 42 8 2009 2009 CD cord-346267-l08ld2cy 42 9 ) ) -RRB- cord-346267-l08ld2cy 42 10 observed observe VBD cord-346267-l08ld2cy 42 11 that that IN cord-346267-l08ld2cy 42 12 synonymous synonymous JJ cord-346267-l08ld2cy 42 13 substitutions substitution NNS cord-346267-l08ld2cy 42 14 appeared appear VBD cord-346267-l08ld2cy 42 15 to to TO cord-346267-l08ld2cy 42 16 saturate saturate VB cord-346267-l08ld2cy 42 17 faster fast RBR cord-346267-l08ld2cy 42 18 than than IN cord-346267-l08ld2cy 42 19 could could MD cord-346267-l08ld2cy 42 20 be be VB cord-346267-l08ld2cy 42 21 handled handle VBN cord-346267-l08ld2cy 42 22 by by IN cord-346267-l08ld2cy 42 23 nucleotide nucleotide JJ cord-346267-l08ld2cy 42 24 models model NNS cord-346267-l08ld2cy 42 25 in in IN cord-346267-l08ld2cy 42 26 mitochondrial mitochondrial JJ cord-346267-l08ld2cy 42 27 DNA dna NN cord-346267-l08ld2cy 42 28 , , , cord-346267-l08ld2cy 42 29 which which WDT cord-346267-l08ld2cy 42 30 is be VBZ cord-346267-l08ld2cy 42 31 subject subject JJ cord-346267-l08ld2cy 42 32 to to IN cord-346267-l08ld2cy 42 33 strong strong JJ cord-346267-l08ld2cy 42 34 overall overall JJ cord-346267-l08ld2cy 42 35 purifying purifying NN cord-346267-l08ld2cy 42 36 selection selection NN cord-346267-l08ld2cy 42 37 . . . cord-346267-l08ld2cy 43 1 Over over IN cord-346267-l08ld2cy 43 2 longer long JJR cord-346267-l08ld2cy 43 3 periods period NNS cord-346267-l08ld2cy 43 4 of of IN cord-346267-l08ld2cy 43 5 evolutionary evolutionary JJ cord-346267-l08ld2cy 43 6 time time NN cord-346267-l08ld2cy 43 7 ( ( -LRB- cord-346267-l08ld2cy 43 8 i.e. i.e. FW cord-346267-l08ld2cy 43 9 , , , cord-346267-l08ld2cy 43 10 along along IN cord-346267-l08ld2cy 43 11 long long JJ cord-346267-l08ld2cy 43 12 internal internal JJ cord-346267-l08ld2cy 43 13 branches branch NNS cord-346267-l08ld2cy 43 14 on on IN cord-346267-l08ld2cy 43 15 a a DT cord-346267-l08ld2cy 43 16 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 43 17 tree tree NN cord-346267-l08ld2cy 43 18 ) ) -RRB- cord-346267-l08ld2cy 43 19 , , , cord-346267-l08ld2cy 43 20 evidence evidence NN cord-346267-l08ld2cy 43 21 of of IN cord-346267-l08ld2cy 43 22 evolution evolution NN cord-346267-l08ld2cy 43 23 at at IN cord-346267-l08ld2cy 43 24 the the DT cord-346267-l08ld2cy 43 25 nucleotide nucleotide JJ cord-346267-l08ld2cy 43 26 level level NN cord-346267-l08ld2cy 43 27 could could MD cord-346267-l08ld2cy 43 28 be be VB cord-346267-l08ld2cy 43 29 lost lose VBN cord-346267-l08ld2cy 43 30 , , , cord-346267-l08ld2cy 43 31 which which WDT cord-346267-l08ld2cy 43 32 could could MD cord-346267-l08ld2cy 43 33 bias bias VB cord-346267-l08ld2cy 43 34 tMRCA tmrca NN cord-346267-l08ld2cy 43 35 estimates estimate NNS cord-346267-l08ld2cy 43 36 towards towards IN cord-346267-l08ld2cy 43 37 younger young JJR cord-346267-l08ld2cy 43 38 dates date NNS cord-346267-l08ld2cy 43 39 ( ( -LRB- cord-346267-l08ld2cy 43 40 i.e. i.e. FW cord-346267-l08ld2cy 43 41 , , , cord-346267-l08ld2cy 43 42 underestimate underestimate VB cord-346267-l08ld2cy 43 43 the the DT cord-346267-l08ld2cy 43 44 length length NN cord-346267-l08ld2cy 43 45 of of IN cord-346267-l08ld2cy 43 46 these these DT cord-346267-l08ld2cy 43 47 branches branch NNS cord-346267-l08ld2cy 43 48 ) ) -RRB- cord-346267-l08ld2cy 43 49 . . . cord-346267-l08ld2cy 44 1 If if IN cord-346267-l08ld2cy 44 2 a a DT cord-346267-l08ld2cy 44 3 substitution substitution NN cord-346267-l08ld2cy 44 4 model model NN cord-346267-l08ld2cy 44 5 can can MD cord-346267-l08ld2cy 44 6 account account VB cord-346267-l08ld2cy 44 7 for for IN cord-346267-l08ld2cy 44 8 this this DT cord-346267-l08ld2cy 44 9 evolution evolution NN cord-346267-l08ld2cy 44 10 , , , cord-346267-l08ld2cy 44 11 we -PRON- PRP cord-346267-l08ld2cy 44 12 hypothesize hypothesize VBP cord-346267-l08ld2cy 44 13 that that IN cord-346267-l08ld2cy 44 14 a a DT cord-346267-l08ld2cy 44 15 comparative comparative JJ cord-346267-l08ld2cy 44 16 analysis analysis NN cord-346267-l08ld2cy 44 17 of of IN cord-346267-l08ld2cy 44 18 recent recent JJ cord-346267-l08ld2cy 44 19 viral viral JJ cord-346267-l08ld2cy 44 20 isolates isolate NNS cord-346267-l08ld2cy 44 21 could could MD cord-346267-l08ld2cy 44 22 be be VB cord-346267-l08ld2cy 44 23 used use VBN cord-346267-l08ld2cy 44 24 to to TO cord-346267-l08ld2cy 44 25 make make VB cord-346267-l08ld2cy 44 26 inference inference NN cord-346267-l08ld2cy 44 27 about about IN cord-346267-l08ld2cy 44 28 ancient ancient JJ cord-346267-l08ld2cy 44 29 viral viral JJ cord-346267-l08ld2cy 44 30 divergence divergence NN cord-346267-l08ld2cy 44 31 events event NNS cord-346267-l08ld2cy 44 32 . . . cord-346267-l08ld2cy 45 1 Our -PRON- PRP$ cord-346267-l08ld2cy 45 2 work work NN cord-346267-l08ld2cy 45 3 addresses address VBZ cord-346267-l08ld2cy 45 4 two two CD cord-346267-l08ld2cy 45 5 questions question NNS cord-346267-l08ld2cy 45 6 . . . cord-346267-l08ld2cy 46 1 First first RB cord-346267-l08ld2cy 46 2 , , , cord-346267-l08ld2cy 46 3 can can MD cord-346267-l08ld2cy 46 4 purifying purify VBG cord-346267-l08ld2cy 46 5 selection selection NN cord-346267-l08ld2cy 46 6 mask mask NN cord-346267-l08ld2cy 46 7 the the DT cord-346267-l08ld2cy 46 8 ancient ancient JJ cord-346267-l08ld2cy 46 9 history history NN cord-346267-l08ld2cy 46 10 of of IN cord-346267-l08ld2cy 46 11 RNA RNA NNP cord-346267-l08ld2cy 46 12 viruses virus NNS cord-346267-l08ld2cy 46 13 ? ? . cord-346267-l08ld2cy 47 1 And and CC cord-346267-l08ld2cy 47 2 second second JJ cord-346267-l08ld2cy 47 3 , , , cord-346267-l08ld2cy 47 4 by by IN cord-346267-l08ld2cy 47 5 using use VBG cord-346267-l08ld2cy 47 6 evolutionary evolutionary JJ cord-346267-l08ld2cy 47 7 models model NNS cord-346267-l08ld2cy 47 8 that that WDT cord-346267-l08ld2cy 47 9 account account VBP cord-346267-l08ld2cy 47 10 for for IN cord-346267-l08ld2cy 47 11 selection selection NN cord-346267-l08ld2cy 47 12 , , , cord-346267-l08ld2cy 47 13 can can MD cord-346267-l08ld2cy 47 14 we -PRON- PRP cord-346267-l08ld2cy 47 15 infer infer VB cord-346267-l08ld2cy 47 16 more more RBR cord-346267-l08ld2cy 47 17 realistic realistic JJ cord-346267-l08ld2cy 47 18 estimates estimate NNS cord-346267-l08ld2cy 47 19 of of IN cord-346267-l08ld2cy 47 20 the the DT cord-346267-l08ld2cy 47 21 ages age NNS cord-346267-l08ld2cy 47 22 of of IN cord-346267-l08ld2cy 47 23 these these DT cord-346267-l08ld2cy 47 24 viruses virus NNS cord-346267-l08ld2cy 47 25 ? ? . cord-346267-l08ld2cy 48 1 We -PRON- PRP cord-346267-l08ld2cy 48 2 investigate investigate VBP cord-346267-l08ld2cy 48 3 these these DT cord-346267-l08ld2cy 48 4 questions question NNS cord-346267-l08ld2cy 48 5 using use VBG cord-346267-l08ld2cy 48 6 two two CD cord-346267-l08ld2cy 48 7 groups group NNS cord-346267-l08ld2cy 48 8 of of IN cord-346267-l08ld2cy 48 9 viruses virus NNS cord-346267-l08ld2cy 48 10 thought think VBN cord-346267-l08ld2cy 48 11 to to TO cord-346267-l08ld2cy 48 12 be be VB cord-346267-l08ld2cy 48 13 older old JJR cord-346267-l08ld2cy 48 14 than than IN cord-346267-l08ld2cy 48 15 current current JJ cord-346267-l08ld2cy 48 16 molecular molecular JJ cord-346267-l08ld2cy 48 17 dating date VBG cord-346267-l08ld2cy 48 18 evidence evidence NN cord-346267-l08ld2cy 48 19 would would MD cord-346267-l08ld2cy 48 20 suggest suggest VB cord-346267-l08ld2cy 48 21 : : : cord-346267-l08ld2cy 48 22 MeV MeV NNP cord-346267-l08ld2cy 48 23 / / SYM cord-346267-l08ld2cy 48 24 RPV RPV NNP cord-346267-l08ld2cy 48 25 / / SYM cord-346267-l08ld2cy 48 26 PPRV PPRV NNP cord-346267-l08ld2cy 48 27 and and CC cord-346267-l08ld2cy 48 28 EBOV EBOV NNP cord-346267-l08ld2cy 48 29 . . . cord-346267-l08ld2cy 49 1 We -PRON- PRP cord-346267-l08ld2cy 49 2 also also RB cord-346267-l08ld2cy 49 3 consider consider VBP cord-346267-l08ld2cy 49 4 avian avian JJ cord-346267-l08ld2cy 49 5 influenza influenza NN cord-346267-l08ld2cy 49 6 virus virus NN cord-346267-l08ld2cy 49 7 ( ( -LRB- cord-346267-l08ld2cy 49 8 AIV)another aiv)another RB cord-346267-l08ld2cy 49 9 well well RB cord-346267-l08ld2cy 49 10 - - HYPH cord-346267-l08ld2cy 49 11 characterized characterize VBN cord-346267-l08ld2cy 49 12 viral viral JJ cord-346267-l08ld2cy 49 13 lineage lineage NN cord-346267-l08ld2cy 49 14 with with IN cord-346267-l08ld2cy 49 15 a a DT cord-346267-l08ld2cy 49 16 young young JJ cord-346267-l08ld2cy 49 17 inferred infer VBN cord-346267-l08ld2cy 49 18 tMRCA tmrca NN cord-346267-l08ld2cy 49 19 and and CC cord-346267-l08ld2cy 49 20 long long JJ cord-346267-l08ld2cy 49 21 internal internal JJ cord-346267-l08ld2cy 49 22 branches branch NNS cord-346267-l08ld2cy 49 23 between between IN cord-346267-l08ld2cy 49 24 different different JJ cord-346267-l08ld2cy 49 25 serotypes serotype NNS cord-346267-l08ld2cy 49 26 Holmes Holmes NNP cord-346267-l08ld2cy 49 27 2006 2006 CD cord-346267-l08ld2cy 49 28 , , , cord-346267-l08ld2cy 49 29 2010 2010 CD cord-346267-l08ld2cy 49 30 ) ) -RRB- cord-346267-l08ld2cy 49 31 . . . cord-346267-l08ld2cy 50 1 The the DT cord-346267-l08ld2cy 50 2 results result NNS cord-346267-l08ld2cy 50 3 presented present VBN cord-346267-l08ld2cy 50 4 here here RB cord-346267-l08ld2cy 50 5 demonstrate demonstrate VBP cord-346267-l08ld2cy 50 6 that that IN cord-346267-l08ld2cy 50 7 not not RB cord-346267-l08ld2cy 50 8 accounting account VBG cord-346267-l08ld2cy 50 9 for for IN cord-346267-l08ld2cy 50 10 purifying purify VBG cord-346267-l08ld2cy 50 11 selection selection NN cord-346267-l08ld2cy 50 12 may may MD cord-346267-l08ld2cy 50 13 bias bias VB cord-346267-l08ld2cy 50 14 tMRCA tmrca NN cord-346267-l08ld2cy 50 15 estimation estimation NN cord-346267-l08ld2cy 50 16 in in IN cord-346267-l08ld2cy 50 17 RNA RNA NNP cord-346267-l08ld2cy 50 18 viruses virus NNS cord-346267-l08ld2cy 50 19 towards towards IN cord-346267-l08ld2cy 50 20 more more RBR cord-346267-l08ld2cy 50 21 recent recent JJ cord-346267-l08ld2cy 50 22 dates date NNS cord-346267-l08ld2cy 50 23 , , , cord-346267-l08ld2cy 50 24 and and CC cord-346267-l08ld2cy 50 25 a a DT cord-346267-l08ld2cy 50 26 degree degree NN cord-346267-l08ld2cy 50 27 of of IN cord-346267-l08ld2cy 50 28 correction correction NN cord-346267-l08ld2cy 50 29 can can MD cord-346267-l08ld2cy 50 30 be be VB cord-346267-l08ld2cy 50 31 realized realize VBN cord-346267-l08ld2cy 50 32 by by IN cord-346267-l08ld2cy 50 33 employing employ VBG cord-346267-l08ld2cy 50 34 more more RBR cord-346267-l08ld2cy 50 35 realistic realistic JJ cord-346267-l08ld2cy 50 36 codon codon NN cord-346267-l08ld2cy 50 37 - - HYPH cord-346267-l08ld2cy 50 38 based base VBN cord-346267-l08ld2cy 50 39 substitution substitution NN cord-346267-l08ld2cy 50 40 models model NNS cord-346267-l08ld2cy 50 41 , , , cord-346267-l08ld2cy 50 42 capable capable JJ cord-346267-l08ld2cy 50 43 of of IN cord-346267-l08ld2cy 50 44 partially partially RB cord-346267-l08ld2cy 50 45 accounting account VBG cord-346267-l08ld2cy 50 46 for for IN cord-346267-l08ld2cy 50 47 the the DT cord-346267-l08ld2cy 50 48 biasing biasing NN cord-346267-l08ld2cy 50 49 effect effect NN cord-346267-l08ld2cy 50 50 of of IN cord-346267-l08ld2cy 50 51 purifying purify VBG cord-346267-l08ld2cy 50 52 selection selection NN cord-346267-l08ld2cy 50 53 . . . cord-346267-l08ld2cy 51 1 For for IN cord-346267-l08ld2cy 51 2 MeV MeV NNP cord-346267-l08ld2cy 51 3 / / SYM cord-346267-l08ld2cy 51 4 RPV RPV NNP cord-346267-l08ld2cy 51 5 / / SYM cord-346267-l08ld2cy 51 6 PPRV PPRV NNP cord-346267-l08ld2cy 51 7 , , , cord-346267-l08ld2cy 51 8 all all DT cord-346267-l08ld2cy 51 9 available available JJ cord-346267-l08ld2cy 51 10 full full JJ cord-346267-l08ld2cy 51 11 - - HYPH cord-346267-l08ld2cy 51 12 length length NN cord-346267-l08ld2cy 51 13 nucleoprotein nucleoprotein JJ cord-346267-l08ld2cy 51 14 sequences sequence NNS cord-346267-l08ld2cy 51 15 with with IN cord-346267-l08ld2cy 51 16 known know VBN cord-346267-l08ld2cy 51 17 years year NNS cord-346267-l08ld2cy 51 18 of of IN cord-346267-l08ld2cy 51 19 isolation isolation NN cord-346267-l08ld2cy 51 20 were be VBD cord-346267-l08ld2cy 51 21 downloaded download VBN cord-346267-l08ld2cy 51 22 from from IN cord-346267-l08ld2cy 51 23 GenBank GenBank NNP cord-346267-l08ld2cy 51 24 . . . cord-346267-l08ld2cy 52 1 Vaccine vaccine NN cord-346267-l08ld2cy 52 2 - - HYPH cord-346267-l08ld2cy 52 3 associated associate VBN cord-346267-l08ld2cy 52 4 sequences sequence NNS cord-346267-l08ld2cy 52 5 and and CC cord-346267-l08ld2cy 52 6 MeV MeV NNP cord-346267-l08ld2cy 52 7 isolates isolate VBZ cord-346267-l08ld2cy 52 8 implicated implicate VBN cord-346267-l08ld2cy 52 9 in in IN cord-346267-l08ld2cy 52 10 subacute subacute JJ cord-346267-l08ld2cy 52 11 sclerosing sclerosing NN cord-346267-l08ld2cy 52 12 panencephalitis panencephalitis NN cord-346267-l08ld2cy 52 13 , , , cord-346267-l08ld2cy 52 14 which which WDT cord-346267-l08ld2cy 52 15 experience experience NN cord-346267-l08ld2cy 52 16 hypermutation hypermutation NN cord-346267-l08ld2cy 53 1 ( ( -LRB- cord-346267-l08ld2cy 53 2 Woelk Woelk NNP cord-346267-l08ld2cy 53 3 et et NNP cord-346267-l08ld2cy 53 4 al al NNP cord-346267-l08ld2cy 53 5 . . . cord-346267-l08ld2cy 54 1 2001 2001 CD cord-346267-l08ld2cy 55 1 ( ( -LRB- cord-346267-l08ld2cy 55 2 Woelk Woelk NNP cord-346267-l08ld2cy 55 3 et et NNP cord-346267-l08ld2cy 55 4 al al NNP cord-346267-l08ld2cy 55 5 . . NNP cord-346267-l08ld2cy 55 6 , , , cord-346267-l08ld2cy 55 7 2002 2002 CD cord-346267-l08ld2cy 55 8 , , , cord-346267-l08ld2cy 55 9 were be VBD cord-346267-l08ld2cy 55 10 excluded exclude VBN cord-346267-l08ld2cy 55 11 . . . cord-346267-l08ld2cy 56 1 The the DT cord-346267-l08ld2cy 56 2 curated curate VBN cord-346267-l08ld2cy 56 3 MeV MeV NNP cord-346267-l08ld2cy 56 4 / / SYM cord-346267-l08ld2cy 56 5 RPV RPV NNP cord-346267-l08ld2cy 56 6 / / SYM cord-346267-l08ld2cy 56 7 PPRV PPRV NNP cord-346267-l08ld2cy 56 8 virus virus NN cord-346267-l08ld2cy 56 9 data datum NNS cord-346267-l08ld2cy 56 10 set set VBN cord-346267-l08ld2cy 56 11 contained contain VBD cord-346267-l08ld2cy 56 12 145 145 CD cord-346267-l08ld2cy 56 13 sequences sequence NNS cord-346267-l08ld2cy 56 14 sampled sample VBN cord-346267-l08ld2cy 56 15 between between IN cord-346267-l08ld2cy 56 16 1981 1981 CD cord-346267-l08ld2cy 56 17 and and CC cord-346267-l08ld2cy 56 18 2007 2007 CD cord-346267-l08ld2cy 56 19 . . . cord-346267-l08ld2cy 57 1 Alignment Alignment NNP cord-346267-l08ld2cy 57 2 was be VBD cord-346267-l08ld2cy 57 3 trivial trivial JJ cord-346267-l08ld2cy 57 4 due due IN cord-346267-l08ld2cy 57 5 to to IN cord-346267-l08ld2cy 57 6 a a DT cord-346267-l08ld2cy 57 7 lack lack NN cord-346267-l08ld2cy 57 8 of of IN cord-346267-l08ld2cy 57 9 insertions insertion NNS cord-346267-l08ld2cy 57 10 or or CC cord-346267-l08ld2cy 57 11 deletions deletion NNS cord-346267-l08ld2cy 57 12 and and CC cord-346267-l08ld2cy 57 13 was be VBD cord-346267-l08ld2cy 57 14 performed perform VBN cord-346267-l08ld2cy 57 15 by by IN cord-346267-l08ld2cy 57 16 eye eye NN cord-346267-l08ld2cy 57 17 . . . cord-346267-l08ld2cy 58 1 Next next RB cord-346267-l08ld2cy 58 2 , , , cord-346267-l08ld2cy 58 3 EBOV EBOV NNP cord-346267-l08ld2cy 58 4 full full JJ cord-346267-l08ld2cy 58 5 - - HYPH cord-346267-l08ld2cy 58 6 length length NN cord-346267-l08ld2cy 58 7 glycoprotein glycoprotein NN cord-346267-l08ld2cy 58 8 sequences sequence NNS cord-346267-l08ld2cy 58 9 with with IN cord-346267-l08ld2cy 58 10 known know VBN cord-346267-l08ld2cy 58 11 years year NNS cord-346267-l08ld2cy 58 12 of of IN cord-346267-l08ld2cy 58 13 isolation isolation NN cord-346267-l08ld2cy 58 14 were be VBD cord-346267-l08ld2cy 58 15 downloaded download VBN cord-346267-l08ld2cy 58 16 from from IN cord-346267-l08ld2cy 58 17 GenBank GenBank NNP cord-346267-l08ld2cy 58 18 . . . cord-346267-l08ld2cy 59 1 Regions region NNS cord-346267-l08ld2cy 59 2 containing contain VBG cord-346267-l08ld2cy 59 3 multiple multiple JJ cord-346267-l08ld2cy 59 4 reading reading NN cord-346267-l08ld2cy 59 5 frames frame NNS cord-346267-l08ld2cy 59 6 ( ( -LRB- cord-346267-l08ld2cy 59 7 Volchkov Volchkov NNP cord-346267-l08ld2cy 59 8 et et NNP cord-346267-l08ld2cy 59 9 al al NNP cord-346267-l08ld2cy 59 10 . . NNP cord-346267-l08ld2cy 59 11 1995 1995 CD cord-346267-l08ld2cy 59 12 ; ; : cord-346267-l08ld2cy 59 13 Sanchez Sanchez NNP cord-346267-l08ld2cy 59 14 et et NNP cord-346267-l08ld2cy 59 15 al al NNP cord-346267-l08ld2cy 59 16 . . . cord-346267-l08ld2cy 59 17 1996 1996 CD cord-346267-l08ld2cy 59 18 ) ) -RRB- cord-346267-l08ld2cy 59 19 and and CC cord-346267-l08ld2cy 59 20 the the DT cord-346267-l08ld2cy 59 21 mucin mucin NN cord-346267-l08ld2cy 59 22 - - HYPH cord-346267-l08ld2cy 59 23 like like JJ cord-346267-l08ld2cy 59 24 region region NN cord-346267-l08ld2cy 59 25 - - , cord-346267-l08ld2cy 59 26 which which WDT cord-346267-l08ld2cy 59 27 evolves evolve VBZ cord-346267-l08ld2cy 59 28 extremely extremely RB cord-346267-l08ld2cy 59 29 rapidly rapidly RB cord-346267-l08ld2cy 59 30 due due JJ cord-346267-l08ld2cy 59 31 to to IN cord-346267-l08ld2cy 59 32 relaxed relaxed JJ cord-346267-l08ld2cy 59 33 selective selective JJ cord-346267-l08ld2cy 59 34 constraints constraint NNS cord-346267-l08ld2cy 59 35 ( ( -LRB- cord-346267-l08ld2cy 59 36 Wertheim Wertheim NNP cord-346267-l08ld2cy 59 37 and and CC cord-346267-l08ld2cy 59 38 Worobey Worobey NNP cord-346267-l08ld2cy 59 39 2009b 2009b CD cord-346267-l08ld2cy 59 40 ) ) -RRB- cord-346267-l08ld2cy 60 1 -were -were RBR cord-346267-l08ld2cy 60 2 removed remove VBN cord-346267-l08ld2cy 60 3 . . . cord-346267-l08ld2cy 61 1 The the DT cord-346267-l08ld2cy 61 2 curated curate VBN cord-346267-l08ld2cy 61 3 EBOV EBOV NNP cord-346267-l08ld2cy 61 4 data datum NNS cord-346267-l08ld2cy 61 5 set set VBN cord-346267-l08ld2cy 61 6 contained contain VBD cord-346267-l08ld2cy 61 7 34 34 CD cord-346267-l08ld2cy 61 8 sequences sequence NNS cord-346267-l08ld2cy 61 9 sampled sample VBN cord-346267-l08ld2cy 61 10 between between IN cord-346267-l08ld2cy 61 11 1976 1976 CD cord-346267-l08ld2cy 61 12 and and CC cord-346267-l08ld2cy 61 13 2005 2005 CD cord-346267-l08ld2cy 61 14 . . . cord-346267-l08ld2cy 62 1 As as IN cord-346267-l08ld2cy 62 2 before before RB cord-346267-l08ld2cy 62 3 , , , cord-346267-l08ld2cy 62 4 alignment alignment NN cord-346267-l08ld2cy 62 5 was be VBD cord-346267-l08ld2cy 62 6 trivial trivial JJ cord-346267-l08ld2cy 62 7 and and CC cord-346267-l08ld2cy 62 8 performed perform VBN cord-346267-l08ld2cy 62 9 by by IN cord-346267-l08ld2cy 62 10 eye eye NN cord-346267-l08ld2cy 62 11 . . . cord-346267-l08ld2cy 63 1 The the DT cord-346267-l08ld2cy 63 2 final final JJ cord-346267-l08ld2cy 63 3 alignment alignment NN cord-346267-l08ld2cy 63 4 , , , cord-346267-l08ld2cy 63 5 AIV AIV NNP cord-346267-l08ld2cy 63 6 neuraminidase neuraminidase NN cord-346267-l08ld2cy 63 7 , , , cord-346267-l08ld2cy 63 8 was be VBD cord-346267-l08ld2cy 63 9 provided provide VBN cord-346267-l08ld2cy 63 10 by by IN cord-346267-l08ld2cy 63 11 Chen Chen NNP cord-346267-l08ld2cy 63 12 and and CC cord-346267-l08ld2cy 63 13 Holmes Holmes NNP cord-346267-l08ld2cy 63 14 ( ( -LRB- cord-346267-l08ld2cy 63 15 2010 2010 CD cord-346267-l08ld2cy 63 16 ) ) -RRB- cord-346267-l08ld2cy 63 17 . . . cord-346267-l08ld2cy 64 1 It -PRON- PRP cord-346267-l08ld2cy 64 2 contained contain VBD cord-346267-l08ld2cy 64 3 270 270 CD cord-346267-l08ld2cy 64 4 sequences sequence NNS cord-346267-l08ld2cy 64 5 sampled sample VBN cord-346267-l08ld2cy 64 6 between between IN cord-346267-l08ld2cy 64 7 1972 1972 CD cord-346267-l08ld2cy 64 8 and and CC cord-346267-l08ld2cy 64 9 2005 2005 CD cord-346267-l08ld2cy 64 10 . . . cord-346267-l08ld2cy 65 1 All all DT cord-346267-l08ld2cy 65 2 three three CD cord-346267-l08ld2cy 65 3 alignments alignment NNS cord-346267-l08ld2cy 65 4 in in IN cord-346267-l08ld2cy 65 5 NEXUS NEXUS NNP cord-346267-l08ld2cy 65 6 format format NN cord-346267-l08ld2cy 65 7 can can MD cord-346267-l08ld2cy 65 8 be be VB cord-346267-l08ld2cy 65 9 downloaded download VBN cord-346267-l08ld2cy 65 10 from from IN cord-346267-l08ld2cy 65 11 http://www.hyphy.org/wiki/codondating http://www.hyphy.org/wiki/codondate VBG cord-346267-l08ld2cy 65 12 . . . cord-346267-l08ld2cy 66 1 Redundant redundant JJ cord-346267-l08ld2cy 66 2 sequences sequence NNS cord-346267-l08ld2cy 66 3 were be VBD cord-346267-l08ld2cy 66 4 excluded exclude VBN cord-346267-l08ld2cy 66 5 from from IN cord-346267-l08ld2cy 66 6 the the DT cord-346267-l08ld2cy 66 7 alignments alignment NNS cord-346267-l08ld2cy 66 8 for for IN cord-346267-l08ld2cy 66 9 branch branch NN cord-346267-l08ld2cy 66 10 length length NN cord-346267-l08ld2cy 66 11 comparisons comparison NNS cord-346267-l08ld2cy 66 12 and and CC cord-346267-l08ld2cy 66 13 selection selection NN cord-346267-l08ld2cy 66 14 analyses analysis NNS cord-346267-l08ld2cy 66 15 . . . cord-346267-l08ld2cy 67 1 We -PRON- PRP cord-346267-l08ld2cy 67 2 used use VBD cord-346267-l08ld2cy 67 3 a a DT cord-346267-l08ld2cy 67 4 Bayesian Bayesian NNP cord-346267-l08ld2cy 67 5 Markov Markov NNP cord-346267-l08ld2cy 67 6 chain chain NN cord-346267-l08ld2cy 67 7 Monte Monte NNP cord-346267-l08ld2cy 67 8 Carlo Carlo NNP cord-346267-l08ld2cy 67 9 ( ( -LRB- cord-346267-l08ld2cy 67 10 BM BM NNP cord-346267-l08ld2cy 67 11 - - HYPH cord-346267-l08ld2cy 67 12 CMC CMC NNP cord-346267-l08ld2cy 67 13 ) ) -RRB- cord-346267-l08ld2cy 67 14 method method NN cord-346267-l08ld2cy 67 15 implemented implement VBN cord-346267-l08ld2cy 67 16 in in IN cord-346267-l08ld2cy 67 17 BEAST BEAST NNP cord-346267-l08ld2cy 67 18 v1.5.4 v1.5.4 NNP cord-346267-l08ld2cy 67 19 for for IN cord-346267-l08ld2cy 67 20 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 67 21 inference inference NN cord-346267-l08ld2cy 67 22 and and CC cord-346267-l08ld2cy 67 23 tMRCA tmrca NN cord-346267-l08ld2cy 67 24 estimation estimation NN cord-346267-l08ld2cy 67 25 . . . cord-346267-l08ld2cy 68 1 For for IN cord-346267-l08ld2cy 68 2 each each DT cord-346267-l08ld2cy 68 3 data data NN cord-346267-l08ld2cy 68 4 set set VBN cord-346267-l08ld2cy 68 5 and and CC cord-346267-l08ld2cy 68 6 substitution substitution NN cord-346267-l08ld2cy 68 7 model model NN cord-346267-l08ld2cy 68 8 , , , cord-346267-l08ld2cy 68 9 four four CD cord-346267-l08ld2cy 68 10 independent independent JJ cord-346267-l08ld2cy 68 11 BMCMC BMCMC NNP cord-346267-l08ld2cy 68 12 runs run VBZ cord-346267-l08ld2cy 68 13 of of IN cord-346267-l08ld2cy 68 14 25 25 CD cord-346267-l08ld2cy 68 15 or or CC cord-346267-l08ld2cy 68 16 50 50 CD cord-346267-l08ld2cy 68 17 million million CD cord-346267-l08ld2cy 68 18 generations generation NNS cord-346267-l08ld2cy 68 19 were be VBD cord-346267-l08ld2cy 68 20 performed perform VBN cord-346267-l08ld2cy 68 21 . . . cord-346267-l08ld2cy 69 1 A a DT cord-346267-l08ld2cy 69 2 codon codon NN cord-346267-l08ld2cy 69 3 - - HYPH cord-346267-l08ld2cy 69 4 based base VBN cord-346267-l08ld2cy 69 5 substitution substitution NN cord-346267-l08ld2cy 69 6 model model NN cord-346267-l08ld2cy 69 7 was be VBD cord-346267-l08ld2cy 69 8 implemented implement VBN cord-346267-l08ld2cy 69 9 in in IN cord-346267-l08ld2cy 69 10 BEAGLE BEAGLE NNP cord-346267-l08ld2cy 69 11 ( ( -LRB- cord-346267-l08ld2cy 69 12 Suchard Suchard NNP cord-346267-l08ld2cy 69 13 and and CC cord-346267-l08ld2cy 69 14 Rambaut Rambaut NNP cord-346267-l08ld2cy 69 15 2009 2009 CD cord-346267-l08ld2cy 69 16 ) ) -RRB- cord-346267-l08ld2cy 69 17 . . . cord-346267-l08ld2cy 70 1 The the DT cord-346267-l08ld2cy 70 2 first first JJ cord-346267-l08ld2cy 70 3 10 10 CD cord-346267-l08ld2cy 70 4 % % NN cord-346267-l08ld2cy 70 5 of of IN cord-346267-l08ld2cy 70 6 the the DT cord-346267-l08ld2cy 70 7 generations generation NNS cord-346267-l08ld2cy 70 8 were be VBD cord-346267-l08ld2cy 70 9 discarded discard VBN cord-346267-l08ld2cy 70 10 as as IN cord-346267-l08ld2cy 70 11 burn burn NN cord-346267-l08ld2cy 70 12 - - HYPH cord-346267-l08ld2cy 70 13 in in NN cord-346267-l08ld2cy 70 14 . . . cord-346267-l08ld2cy 71 1 All all DT cord-346267-l08ld2cy 71 2 BMCMC BMCMC NNP cord-346267-l08ld2cy 71 3 analyses analysis NNS cord-346267-l08ld2cy 71 4 were be VBD cord-346267-l08ld2cy 71 5 performed perform VBN cord-346267-l08ld2cy 71 6 using use VBG cord-346267-l08ld2cy 71 7 an an DT cord-346267-l08ld2cy 71 8 uncorrelated uncorrelated JJ cord-346267-l08ld2cy 71 9 lognormal lognormal JJ cord-346267-l08ld2cy 71 10 relaxed relaxed JJ cord-346267-l08ld2cy 71 11 molecular molecular JJ cord-346267-l08ld2cy 71 12 clock clock NN cord-346267-l08ld2cy 71 13 and and CC cord-346267-l08ld2cy 71 14 a a DT cord-346267-l08ld2cy 71 15 Bayesian bayesian JJ cord-346267-l08ld2cy 71 16 skyline skyline NN cord-346267-l08ld2cy 71 17 plot plot NN cord-346267-l08ld2cy 71 18 coalescent coalescent NN cord-346267-l08ld2cy 71 19 prior prior RB cord-346267-l08ld2cy 71 20 , , , cord-346267-l08ld2cy 71 21 which which WDT cord-346267-l08ld2cy 71 22 places place VBZ cord-346267-l08ld2cy 71 23 the the DT cord-346267-l08ld2cy 71 24 fewest few JJS cord-346267-l08ld2cy 71 25 demographic demographic JJ cord-346267-l08ld2cy 71 26 constraints constraint NNS cord-346267-l08ld2cy 71 27 on on IN cord-346267-l08ld2cy 71 28 the the DT cord-346267-l08ld2cy 71 29 analysis analysis NN cord-346267-l08ld2cy 71 30 . . . cord-346267-l08ld2cy 72 1 Tracer tracer NN cord-346267-l08ld2cy 72 2 v1.5 v1.5 NN cord-346267-l08ld2cy 72 3 was be VBD cord-346267-l08ld2cy 72 4 used use VBN cord-346267-l08ld2cy 72 5 to to TO cord-346267-l08ld2cy 72 6 check check VB cord-346267-l08ld2cy 72 7 for for IN cord-346267-l08ld2cy 72 8 convergence convergence NN cord-346267-l08ld2cy 72 9 and and CC cord-346267-l08ld2cy 72 10 adequate adequate JJ cord-346267-l08ld2cy 72 11 mixing mixing NN cord-346267-l08ld2cy 72 12 ( ( -LRB- cord-346267-l08ld2cy 72 13 i.e. i.e. FW cord-346267-l08ld2cy 72 14 , , , cord-346267-l08ld2cy 72 15 estimated estimate VBN cord-346267-l08ld2cy 72 16 sample sample NN cord-346267-l08ld2cy 72 17 size size NN cord-346267-l08ld2cy 72 18 > > XX cord-346267-l08ld2cy 73 1 200 200 CD cord-346267-l08ld2cy 73 2 for for IN cord-346267-l08ld2cy 73 3 all all DT cord-346267-l08ld2cy 73 4 relevant relevant JJ cord-346267-l08ld2cy 73 5 parameters parameter NNS cord-346267-l08ld2cy 73 6 ) ) -RRB- cord-346267-l08ld2cy 73 7 . . . cord-346267-l08ld2cy 74 1 Finally finally RB cord-346267-l08ld2cy 74 2 , , , cord-346267-l08ld2cy 74 3 the the DT cord-346267-l08ld2cy 74 4 maximum maximum JJ cord-346267-l08ld2cy 74 5 clade clade NN cord-346267-l08ld2cy 74 6 credibility credibility NN cord-346267-l08ld2cy 74 7 ( ( -LRB- cord-346267-l08ld2cy 74 8 MCC MCC NNP cord-346267-l08ld2cy 74 9 ) ) -RRB- cord-346267-l08ld2cy 74 10 phylogeny phylogeny NN cord-346267-l08ld2cy 74 11 was be VBD cord-346267-l08ld2cy 74 12 identified identify VBN cord-346267-l08ld2cy 74 13 and and CC cord-346267-l08ld2cy 74 14 annotated annotate VBN cord-346267-l08ld2cy 74 15 using use VBG cord-346267-l08ld2cy 74 16 the the DT cord-346267-l08ld2cy 74 17 posterior posterior JJ cord-346267-l08ld2cy 74 18 distribution distribution NN cord-346267-l08ld2cy 74 19 of of IN cord-346267-l08ld2cy 74 20 trees tree NNS cord-346267-l08ld2cy 74 21 . . . cord-346267-l08ld2cy 75 1 Substitution substitution NN cord-346267-l08ld2cy 75 2 rates rate NNS cord-346267-l08ld2cy 75 3 and and CC cord-346267-l08ld2cy 75 4 tMRCAs tmrcas CD cord-346267-l08ld2cy 75 5 are be VBP cord-346267-l08ld2cy 75 6 reported report VBN cord-346267-l08ld2cy 75 7 as as IN cord-346267-l08ld2cy 75 8 mean mean JJ cord-346267-l08ld2cy 75 9 and and CC cord-346267-l08ld2cy 75 10 95 95 CD cord-346267-l08ld2cy 75 11 % % NN cord-346267-l08ld2cy 75 12 highest high JJS cord-346267-l08ld2cy 75 13 posterior posterior JJ cord-346267-l08ld2cy 75 14 density density NN cord-346267-l08ld2cy 75 15 values value NNS cord-346267-l08ld2cy 75 16 . . . cord-346267-l08ld2cy 76 1 An an DT cord-346267-l08ld2cy 76 2 earlier early JJR cord-346267-l08ld2cy 76 3 study study NN cord-346267-l08ld2cy 76 4 ) ) -RRB- cord-346267-l08ld2cy 76 5 developed develop VBD cord-346267-l08ld2cy 76 6 a a DT cord-346267-l08ld2cy 76 7 hierarchy hierarchy NN cord-346267-l08ld2cy 76 8 of of IN cord-346267-l08ld2cy 76 9 five five CD cord-346267-l08ld2cy 76 10 models model NNS cord-346267-l08ld2cy 76 11 - - : cord-346267-l08ld2cy 76 12 all all DT cord-346267-l08ld2cy 76 13 extensions extension NNS cord-346267-l08ld2cy 76 14 of of IN cord-346267-l08ld2cy 76 15 the the DT cord-346267-l08ld2cy 76 16 Muse Muse NNP cord-346267-l08ld2cy 76 17 - - HYPH cord-346267-l08ld2cy 76 18 Gaut Gaut NNP cord-346267-l08ld2cy 76 19 probabilistic probabilistic JJ cord-346267-l08ld2cy 76 20 model model NN cord-346267-l08ld2cy 76 21 of of IN cord-346267-l08ld2cy 76 22 sequence sequence NN cord-346267-l08ld2cy 76 23 evolution evolution NN cord-346267-l08ld2cy 76 24 ( ( -LRB- cord-346267-l08ld2cy 76 25 Muse Muse NNP cord-346267-l08ld2cy 76 26 and and CC cord-346267-l08ld2cy 76 27 Gaut Gaut NNP cord-346267-l08ld2cy 76 28 1994 1994 CD cord-346267-l08ld2cy 76 29 ) ) -RRB- cord-346267-l08ld2cy 77 1 -of -of : cord-346267-l08ld2cy 77 2 differing differ VBG cord-346267-l08ld2cy 77 3 complexities complexity NNS cord-346267-l08ld2cy 77 4 that that WDT cord-346267-l08ld2cy 77 5 incorporate incorporate VBP cord-346267-l08ld2cy 77 6 site site NN cord-346267-l08ld2cy 77 7 - - HYPH cord-346267-l08ld2cy 77 8 tosite tosite JJ cord-346267-l08ld2cy 77 9 and and CC cord-346267-l08ld2cy 77 10 lineage lineage NN cord-346267-l08ld2cy 77 11 - - HYPH cord-346267-l08ld2cy 77 12 to to IN cord-346267-l08ld2cy 77 13 - - HYPH cord-346267-l08ld2cy 77 14 lineage lineage NN cord-346267-l08ld2cy 77 15 variation variation NN cord-346267-l08ld2cy 77 16 of of IN cord-346267-l08ld2cy 77 17 synonymous synonymous JJ cord-346267-l08ld2cy 77 18 ( ( -LRB- cord-346267-l08ld2cy 77 19 α α NN cord-346267-l08ld2cy 77 20 ) ) -RRB- cord-346267-l08ld2cy 77 21 and and CC cord-346267-l08ld2cy 77 22 nonsynonymous nonsynonymous NNP cord-346267-l08ld2cy 77 23 ( ( -LRB- cord-346267-l08ld2cy 77 24 β β NN cord-346267-l08ld2cy 77 25 ) ) -RRB- cord-346267-l08ld2cy 77 26 substitution substitution NN cord-346267-l08ld2cy 77 27 rates rate NNS cord-346267-l08ld2cy 77 28 . . . cord-346267-l08ld2cy 78 1 We -PRON- PRP cord-346267-l08ld2cy 78 2 investigated investigate VBD cord-346267-l08ld2cy 78 3 the the DT cord-346267-l08ld2cy 78 4 ability ability NN cord-346267-l08ld2cy 78 5 of of IN cord-346267-l08ld2cy 78 6 these these DT cord-346267-l08ld2cy 78 7 models model NNS cord-346267-l08ld2cy 78 8 to to TO cord-346267-l08ld2cy 78 9 accurately accurately RB cord-346267-l08ld2cy 78 10 infer infer VB cord-346267-l08ld2cy 78 11 branch branch NN cord-346267-l08ld2cy 78 12 lengths length NNS cord-346267-l08ld2cy 78 13 on on IN cord-346267-l08ld2cy 78 14 our -PRON- PRP$ cord-346267-l08ld2cy 78 15 viral viral JJ cord-346267-l08ld2cy 78 16 data data NN cord-346267-l08ld2cy 78 17 sets set NNS cord-346267-l08ld2cy 78 18 . . . cord-346267-l08ld2cy 79 1 The the DT cord-346267-l08ld2cy 79 2 five five CD cord-346267-l08ld2cy 79 3 models model NNS cord-346267-l08ld2cy 79 4 are be VBP cord-346267-l08ld2cy 79 5 summarized summarize VBN cord-346267-l08ld2cy 79 6 below below IN cord-346267-l08ld2cy 79 7 , , , cord-346267-l08ld2cy 79 8 with with IN cord-346267-l08ld2cy 79 9 full full JJ cord-346267-l08ld2cy 79 10 details detail NNS cord-346267-l08ld2cy 79 11 available available JJ cord-346267-l08ld2cy 79 12 in in IN cord-346267-l08ld2cy 79 13 the the DT cord-346267-l08ld2cy 79 14 original original JJ cord-346267-l08ld2cy 79 15 manuscript manuscript NN cord-346267-l08ld2cy 79 16 . . . cord-346267-l08ld2cy 80 1 Note note VB cord-346267-l08ld2cy 80 2 that that IN cord-346267-l08ld2cy 80 3 in in IN cord-346267-l08ld2cy 80 4 all all DT cord-346267-l08ld2cy 80 5 models model NNS cord-346267-l08ld2cy 80 6 , , , cord-346267-l08ld2cy 80 7 E E NNS cord-346267-l08ld2cy 80 8 [ [ -LRB- cord-346267-l08ld2cy 80 9 α α NN cord-346267-l08ld2cy 80 10 ] ] -RRB- cord-346267-l08ld2cy 80 11 = = SYM cord-346267-l08ld2cy 80 12 1 1 CD cord-346267-l08ld2cy 80 13 to to TO cord-346267-l08ld2cy 80 14 ensure ensure VB cord-346267-l08ld2cy 80 15 identifiability identifiability NN cord-346267-l08ld2cy 80 16 . . . cord-346267-l08ld2cy 81 1 Constant constant JJ cord-346267-l08ld2cy 81 2 Rates rate NNS cord-346267-l08ld2cy 81 3 : : : cord-346267-l08ld2cy 81 4 The the DT cord-346267-l08ld2cy 81 5 baseline baseline JJ cord-346267-l08ld2cy 81 6 model model NN cord-346267-l08ld2cy 81 7 , , , cord-346267-l08ld2cy 81 8 which which WDT cord-346267-l08ld2cy 81 9 extends extend VBZ cord-346267-l08ld2cy 81 10 the the DT cord-346267-l08ld2cy 81 11 original original JJ cord-346267-l08ld2cy 81 12 Muse Muse NNP cord-346267-l08ld2cy 81 13 - - HYPH cord-346267-l08ld2cy 81 14 Gaut Gaut NNP cord-346267-l08ld2cy 81 15 evolutionary evolutionary JJ cord-346267-l08ld2cy 81 16 model model NN cord-346267-l08ld2cy 81 17 by by IN cord-346267-l08ld2cy 81 18 incorporating incorporate VBG cord-346267-l08ld2cy 81 19 general general JJ cord-346267-l08ld2cy 81 20 nucleotide nucleotide JJ cord-346267-l08ld2cy 81 21 substitution substitution NN cord-346267-l08ld2cy 81 22 biases bias NNS cord-346267-l08ld2cy 81 23 ( ( -LRB- cord-346267-l08ld2cy 81 24 MG94 MG94 NNP cord-346267-l08ld2cy 81 25 × × NNP cord-346267-l08ld2cy 81 26 REV REV NNP cord-346267-l08ld2cy 81 27 ) ) -RRB- cord-346267-l08ld2cy 81 28 . . . cord-346267-l08ld2cy 82 1 α α NNP cord-346267-l08ld2cy 82 2 and and CC cord-346267-l08ld2cy 82 3 β β NN cord-346267-l08ld2cy 82 4 rates rate NNS cord-346267-l08ld2cy 82 5 are be VBP cord-346267-l08ld2cy 82 6 constant constant JJ cord-346267-l08ld2cy 82 7 across across IN cord-346267-l08ld2cy 82 8 sites site NNS cord-346267-l08ld2cy 82 9 and and CC cord-346267-l08ld2cy 82 10 lineages lineage NNS cord-346267-l08ld2cy 82 11 . . . cord-346267-l08ld2cy 83 1 Proportional proportional JJ cord-346267-l08ld2cy 83 2 : : : cord-346267-l08ld2cy 84 1 A a DT cord-346267-l08ld2cy 84 2 direct direct JJ cord-346267-l08ld2cy 84 3 analog analog NN cord-346267-l08ld2cy 84 4 to to TO cord-346267-l08ld2cy 84 5 nucleotide nucleotide VB cord-346267-l08ld2cy 84 6 + + CC cord-346267-l08ld2cy 84 7 Γ γ NN cord-346267-l08ld2cy 84 8 4 4 CD cord-346267-l08ld2cy 84 9 models model NNS cord-346267-l08ld2cy 84 10 ; ; : cord-346267-l08ld2cy 84 11 β β NN cord-346267-l08ld2cy 84 12 = = SYM cord-346267-l08ld2cy 84 13 ωα ωα NNP cord-346267-l08ld2cy 84 14 and and CC cord-346267-l08ld2cy 84 15 α α NN cord-346267-l08ld2cy 84 16 varies vary VBZ cord-346267-l08ld2cy 84 17 from from IN cord-346267-l08ld2cy 84 18 site site NN cord-346267-l08ld2cy 84 19 to to IN cord-346267-l08ld2cy 84 20 site site NN cord-346267-l08ld2cy 84 21 according accord VBG cord-346267-l08ld2cy 84 22 to to IN cord-346267-l08ld2cy 84 23 a a DT cord-346267-l08ld2cy 84 24 three three CD cord-346267-l08ld2cy 84 25 - - HYPH cord-346267-l08ld2cy 84 26 bin bin NN cord-346267-l08ld2cy 84 27 general general JJ cord-346267-l08ld2cy 84 28 discrete discrete JJ cord-346267-l08ld2cy 84 29 distribution distribution NN cord-346267-l08ld2cy 84 30 ( ( -LRB- cord-346267-l08ld2cy 84 31 GDD GDD NNP cord-346267-l08ld2cy 84 32 ) ) -RRB- cord-346267-l08ld2cy 84 33 . . . cord-346267-l08ld2cy 85 1 Nonsynonymous Nonsynonymous NNP cord-346267-l08ld2cy 85 2 : : : cord-346267-l08ld2cy 86 1 A a DT cord-346267-l08ld2cy 86 2 standard standard JJ cord-346267-l08ld2cy 86 3 " " `` cord-346267-l08ld2cy 86 4 selection selection NN cord-346267-l08ld2cy 86 5 " " '' cord-346267-l08ld2cy 86 6 model model NN cord-346267-l08ld2cy 86 7 , , , cord-346267-l08ld2cy 86 8 where where WRB cord-346267-l08ld2cy 86 9 α α NN cord-346267-l08ld2cy 86 10 rates rate NNS cord-346267-l08ld2cy 86 11 are be VBP cord-346267-l08ld2cy 86 12 constant constant JJ cord-346267-l08ld2cy 86 13 and and CC cord-346267-l08ld2cy 86 14 β β NN cord-346267-l08ld2cy 86 15 rates rate NNS cord-346267-l08ld2cy 86 16 are be VBP cord-346267-l08ld2cy 86 17 drawn draw VBN cord-346267-l08ld2cy 86 18 from from IN cord-346267-l08ld2cy 86 19 a a DT cord-346267-l08ld2cy 86 20 3bin 3bin CD cord-346267-l08ld2cy 86 21 GDD GDD NNP cord-346267-l08ld2cy 86 22 . . . cord-346267-l08ld2cy 87 1 Dual dual JJ cord-346267-l08ld2cy 87 2 : : : cord-346267-l08ld2cy 88 1 A a DT cord-346267-l08ld2cy 88 2 model model NN cord-346267-l08ld2cy 88 3 , , , cord-346267-l08ld2cy 88 4 where where WRB cord-346267-l08ld2cy 88 5 both both CC cord-346267-l08ld2cy 88 6 α α NN cord-346267-l08ld2cy 88 7 and and CC cord-346267-l08ld2cy 88 8 β β NN cord-346267-l08ld2cy 88 9 rates rate NNS cord-346267-l08ld2cy 88 10 vary vary VBP cord-346267-l08ld2cy 88 11 from from IN cord-346267-l08ld2cy 88 12 site site NN cord-346267-l08ld2cy 88 13 to to IN cord-346267-l08ld2cy 88 14 site site NN cord-346267-l08ld2cy 88 15 , , , cord-346267-l08ld2cy 88 16 based base VBN cord-346267-l08ld2cy 88 17 on on IN cord-346267-l08ld2cy 88 18 independent independent JJ cord-346267-l08ld2cy 88 19 three three CD cord-346267-l08ld2cy 88 20 - - HYPH cord-346267-l08ld2cy 88 21 bin bin NN cord-346267-l08ld2cy 88 22 GDD GDD NNP cord-346267-l08ld2cy 88 23 distributions distribution NNS cord-346267-l08ld2cy 88 24 . . . cord-346267-l08ld2cy 89 1 Lineage+Dual lineage+dual CC cord-346267-l08ld2cy 89 2 : : : cord-346267-l08ld2cy 89 3 In in IN cord-346267-l08ld2cy 89 4 addition addition NN cord-346267-l08ld2cy 89 5 to to IN cord-346267-l08ld2cy 89 6 site site NN cord-346267-l08ld2cy 89 7 - - HYPH cord-346267-l08ld2cy 89 8 to to IN cord-346267-l08ld2cy 89 9 - - HYPH cord-346267-l08ld2cy 89 10 site site NN cord-346267-l08ld2cy 89 11 rate rate NN cord-346267-l08ld2cy 89 12 variation variation NN cord-346267-l08ld2cy 89 13 in in IN cord-346267-l08ld2cy 89 14 α α NNP cord-346267-l08ld2cy 89 15 and and CC cord-346267-l08ld2cy 89 16 β β NN cord-346267-l08ld2cy 89 17 , , , cord-346267-l08ld2cy 89 18 some some DT cord-346267-l08ld2cy 89 19 ( ( -LRB- cord-346267-l08ld2cy 89 20 or or CC cord-346267-l08ld2cy 89 21 all all DT cord-346267-l08ld2cy 89 22 ) ) -RRB- cord-346267-l08ld2cy 89 23 lineages lineage NNS cord-346267-l08ld2cy 89 24 are be VBP cord-346267-l08ld2cy 89 25 endowed endow VBN cord-346267-l08ld2cy 89 26 with with IN cord-346267-l08ld2cy 89 27 their -PRON- PRP$ cord-346267-l08ld2cy 89 28 own own JJ cord-346267-l08ld2cy 89 29 mean mean NN cord-346267-l08ld2cy 89 30 E e NN cord-346267-l08ld2cy 89 31 [ [ -LRB- cord-346267-l08ld2cy 89 32 β]/E β]/e XX cord-346267-l08ld2cy 89 33 [ [ -LRB- cord-346267-l08ld2cy 89 34 α α NN cord-346267-l08ld2cy 89 35 ] ] -RRB- cord-346267-l08ld2cy 89 36 ratios ratio NNS cord-346267-l08ld2cy 89 37 , , , cord-346267-l08ld2cy 89 38 to to TO cord-346267-l08ld2cy 89 39 correct correct VB cord-346267-l08ld2cy 89 40 for for IN cord-346267-l08ld2cy 89 41 " " `` cord-346267-l08ld2cy 89 42 lineagespecific lineagespecific JJ cord-346267-l08ld2cy 89 43 " " '' cord-346267-l08ld2cy 89 44 effects effect NNS cord-346267-l08ld2cy 89 45 of of IN cord-346267-l08ld2cy 89 46 selection selection NN cord-346267-l08ld2cy 89 47 . . . cord-346267-l08ld2cy 90 1 Unlike unlike IN cord-346267-l08ld2cy 90 2 the the DT cord-346267-l08ld2cy 90 3 original original JJ cord-346267-l08ld2cy 90 4 implementation implementation NN cord-346267-l08ld2cy 90 5 , , , cord-346267-l08ld2cy 90 6 our -PRON- PRP$ cord-346267-l08ld2cy 90 7 version version NN cord-346267-l08ld2cy 90 8 of of IN cord-346267-l08ld2cy 90 9 the the DT cord-346267-l08ld2cy 90 10 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 90 11 model model NN cord-346267-l08ld2cy 90 12 treated treat VBD cord-346267-l08ld2cy 90 13 certain certain JJ cord-346267-l08ld2cy 90 14 interior interior JJ cord-346267-l08ld2cy 90 15 lineages lineage NNS cord-346267-l08ld2cy 90 16 differently differently RB cord-346267-l08ld2cy 90 17 to to TO cord-346267-l08ld2cy 90 18 better better RB cord-346267-l08ld2cy 90 19 reflect reflect VB cord-346267-l08ld2cy 90 20 the the DT cord-346267-l08ld2cy 90 21 biological biological JJ cord-346267-l08ld2cy 90 22 reality reality NN cord-346267-l08ld2cy 90 23 of of IN cord-346267-l08ld2cy 90 24 the the DT cord-346267-l08ld2cy 90 25 sample sample NN cord-346267-l08ld2cy 90 26 . . . cord-346267-l08ld2cy 91 1 Branches branch NNS cord-346267-l08ld2cy 91 2 which which WDT cord-346267-l08ld2cy 91 3 separated separate VBD cord-346267-l08ld2cy 91 4 clades clade NNS cord-346267-l08ld2cy 91 5 containing contain VBG cord-346267-l08ld2cy 91 6 different different JJ cord-346267-l08ld2cy 91 7 viral viral JJ cord-346267-l08ld2cy 91 8 species specie NNS cord-346267-l08ld2cy 91 9 ( ( -LRB- cord-346267-l08ld2cy 91 10 MeV MeV NNP cord-346267-l08ld2cy 91 11 / / SYM cord-346267-l08ld2cy 91 12 RPV RPV NNP cord-346267-l08ld2cy 91 13 / / SYM cord-346267-l08ld2cy 91 14 PPRV PPRV NNP cord-346267-l08ld2cy 91 15 ) ) -RRB- cord-346267-l08ld2cy 91 16 , , , cord-346267-l08ld2cy 91 17 subtypes subtype NNS cord-346267-l08ld2cy 91 18 ( ( -LRB- cord-346267-l08ld2cy 91 19 EBOV EBOV NNP cord-346267-l08ld2cy 91 20 ) ) -RRB- cord-346267-l08ld2cy 91 21 , , , cord-346267-l08ld2cy 91 22 and and CC cord-346267-l08ld2cy 91 23 serotypes serotype NNS cord-346267-l08ld2cy 91 24 ( ( -LRB- cord-346267-l08ld2cy 91 25 AIV AIV NNP cord-346267-l08ld2cy 91 26 ) ) -RRB- cord-346267-l08ld2cy 91 27 were be VBD cord-346267-l08ld2cy 91 28 assigned assign VBN cord-346267-l08ld2cy 91 29 separate separate JJ cord-346267-l08ld2cy 91 30 E e NN cord-346267-l08ld2cy 91 31 [ [ -LRB- cord-346267-l08ld2cy 91 32 β]/E β]/e XX cord-346267-l08ld2cy 91 33 [ [ -LRB- cord-346267-l08ld2cy 91 34 α α NN cord-346267-l08ld2cy 91 35 ] ] -RRB- cord-346267-l08ld2cy 91 36 ratio ratio NN cord-346267-l08ld2cy 91 37 parameters parameter NNS cord-346267-l08ld2cy 91 38 because because IN cord-346267-l08ld2cy 91 39 they -PRON- PRP cord-346267-l08ld2cy 91 40 represent represent VBP cord-346267-l08ld2cy 91 41 different different JJ cord-346267-l08ld2cy 91 42 timescales timescale NNS cord-346267-l08ld2cy 91 43 and and CC cord-346267-l08ld2cy 91 44 selective selective JJ cord-346267-l08ld2cy 91 45 regimes regime NNS cord-346267-l08ld2cy 91 46 compared compare VBN cord-346267-l08ld2cy 91 47 with with IN cord-346267-l08ld2cy 91 48 terminal terminal JJ cord-346267-l08ld2cy 91 49 branches branch NNS cord-346267-l08ld2cy 91 50 . . . cord-346267-l08ld2cy 92 1 Longer long JJR cord-346267-l08ld2cy 92 2 internal internal JJ cord-346267-l08ld2cy 92 3 branches branch NNS cord-346267-l08ld2cy 92 4 are be VBP cord-346267-l08ld2cy 92 5 expected expect VBN cord-346267-l08ld2cy 92 6 to to TO cord-346267-l08ld2cy 92 7 bear bear VB cord-346267-l08ld2cy 92 8 the the DT cord-346267-l08ld2cy 92 9 mark mark NN cord-346267-l08ld2cy 92 10 of of IN cord-346267-l08ld2cy 92 11 stronger strong JJR cord-346267-l08ld2cy 92 12 purifying purifying NN cord-346267-l08ld2cy 92 13 selection selection NN cord-346267-l08ld2cy 93 1 ( ( -LRB- cord-346267-l08ld2cy 93 2 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 93 3 et et FW cord-346267-l08ld2cy 93 4 al al NNP cord-346267-l08ld2cy 93 5 . . . cord-346267-l08ld2cy 94 1 2006 2006 CD cord-346267-l08ld2cy 94 2 ; ; : cord-346267-l08ld2cy 94 3 Pybus Pybus NNP cord-346267-l08ld2cy 94 4 et et NNP cord-346267-l08ld2cy 94 5 al al NNP cord-346267-l08ld2cy 94 6 . . NNP cord-346267-l08ld2cy 94 7 2007 2007 CD cord-346267-l08ld2cy 94 8 ) ) -RRB- cord-346267-l08ld2cy 94 9 . . . cord-346267-l08ld2cy 95 1 We -PRON- PRP cord-346267-l08ld2cy 95 2 considered consider VBD cord-346267-l08ld2cy 95 3 two two CD cord-346267-l08ld2cy 95 4 variations variation NNS cord-346267-l08ld2cy 95 5 of of IN cord-346267-l08ld2cy 95 6 this this DT cord-346267-l08ld2cy 95 7 model model NN cord-346267-l08ld2cy 95 8 : : : cord-346267-l08ld2cy 95 9 Either either CC cord-346267-l08ld2cy 95 10 1 1 LS cord-346267-l08ld2cy 95 11 ) ) -RRB- cord-346267-l08ld2cy 95 12 all all DT cord-346267-l08ld2cy 95 13 long long JJ cord-346267-l08ld2cy 95 14 internal internal JJ cord-346267-l08ld2cy 95 15 lineages lineage NNS cord-346267-l08ld2cy 95 16 share share VBP cord-346267-l08ld2cy 95 17 the the DT cord-346267-l08ld2cy 95 18 same same JJ cord-346267-l08ld2cy 95 19 ratio ratio NN cord-346267-l08ld2cy 95 20 parameter parameter NN cord-346267-l08ld2cy 95 21 , , , cord-346267-l08ld2cy 95 22 and and CC cord-346267-l08ld2cy 95 23 the the DT cord-346267-l08ld2cy 95 24 remaining remain VBG cord-346267-l08ld2cy 95 25 branches branch NNS cord-346267-l08ld2cy 95 26 share share VBP cord-346267-l08ld2cy 95 27 another another DT cord-346267-l08ld2cy 95 28 global global JJ cord-346267-l08ld2cy 95 29 ratio ratio NN cord-346267-l08ld2cy 95 30 parameter parameter NN cord-346267-l08ld2cy 95 31 ( ( -LRB- cord-346267-l08ld2cy 95 32 two two CD cord-346267-l08ld2cy 95 33 - - HYPH cord-346267-l08ld2cy 95 34 rate rate NN cord-346267-l08ld2cy 95 35 model model NN cord-346267-l08ld2cy 95 36 ) ) -RRB- cord-346267-l08ld2cy 95 37 or or CC cord-346267-l08ld2cy 95 38 2 2 CD cord-346267-l08ld2cy 95 39 ) ) -RRB- cord-346267-l08ld2cy 96 1 each each DT cord-346267-l08ld2cy 96 2 deep deep JJ cord-346267-l08ld2cy 96 3 lineage lineage NN cord-346267-l08ld2cy 96 4 possesses possess VBZ cord-346267-l08ld2cy 96 5 its -PRON- PRP$ cord-346267-l08ld2cy 96 6 own own JJ cord-346267-l08ld2cy 96 7 ratio ratio NN cord-346267-l08ld2cy 96 8 parameter parameter NN cord-346267-l08ld2cy 96 9 , , , cord-346267-l08ld2cy 96 10 and and CC cord-346267-l08ld2cy 96 11 the the DT cord-346267-l08ld2cy 96 12 remaining remain VBG cord-346267-l08ld2cy 96 13 branches branch NNS cord-346267-l08ld2cy 96 14 share share VBP cord-346267-l08ld2cy 96 15 a a DT cord-346267-l08ld2cy 96 16 global global JJ cord-346267-l08ld2cy 96 17 ratio ratio NN cord-346267-l08ld2cy 96 18 parameter parameter NN cord-346267-l08ld2cy 96 19 ( ( -LRB- cord-346267-l08ld2cy 96 20 multirate multirate NNP cord-346267-l08ld2cy 96 21 model model NN cord-346267-l08ld2cy 96 22 ) ) -RRB- cord-346267-l08ld2cy 96 23 . . . cord-346267-l08ld2cy 97 1 For for IN cord-346267-l08ld2cy 97 2 each each DT cord-346267-l08ld2cy 97 3 data data NN cord-346267-l08ld2cy 97 4 set set VBN cord-346267-l08ld2cy 97 5 , , , cord-346267-l08ld2cy 97 6 we -PRON- PRP cord-346267-l08ld2cy 97 7 selected select VBD cord-346267-l08ld2cy 97 8 the the DT cord-346267-l08ld2cy 97 9 model model NN cord-346267-l08ld2cy 97 10 with with IN cord-346267-l08ld2cy 97 11 the the DT cord-346267-l08ld2cy 97 12 best good JJS cord-346267-l08ld2cy 97 13 Akaike Akaike NNP cord-346267-l08ld2cy 97 14 information information NN cord-346267-l08ld2cy 97 15 criterion criterion NN cord-346267-l08ld2cy 97 16 ( ( -LRB- cord-346267-l08ld2cy 97 17 AIC AIC NNP cord-346267-l08ld2cy 97 18 ) ) -RRB- cord-346267-l08ld2cy 97 19 score score NN cord-346267-l08ld2cy 97 20 for for IN cord-346267-l08ld2cy 97 21 further further JJ cord-346267-l08ld2cy 97 22 analysis analysis NN cord-346267-l08ld2cy 97 23 . . . cord-346267-l08ld2cy 98 1 Note note VB cord-346267-l08ld2cy 98 2 that that IN cord-346267-l08ld2cy 98 3 we -PRON- PRP cord-346267-l08ld2cy 98 4 use use VBP cord-346267-l08ld2cy 98 5 the the DT cord-346267-l08ld2cy 98 6 ratio ratio NN cord-346267-l08ld2cy 98 7 of of IN cord-346267-l08ld2cy 98 8 the the DT cord-346267-l08ld2cy 98 9 means mean NNS cord-346267-l08ld2cy 98 10 E e NN cord-346267-l08ld2cy 98 11 is be VBZ cord-346267-l08ld2cy 98 12 possible possible JJ cord-346267-l08ld2cy 98 13 under under IN cord-346267-l08ld2cy 98 14 Dual Dual NNP cord-346267-l08ld2cy 98 15 and and CC cord-346267-l08ld2cy 98 16 Lin Lin NNP cord-346267-l08ld2cy 98 17 - - HYPH cord-346267-l08ld2cy 98 18 eage+Dual eage+Dual NNP cord-346267-l08ld2cy 98 19 models model NNS cord-346267-l08ld2cy 98 20 , , , cord-346267-l08ld2cy 98 21 rendering render VBG cord-346267-l08ld2cy 98 22 the the DT cord-346267-l08ld2cy 98 23 mean mean NN cord-346267-l08ld2cy 98 24 of of IN cord-346267-l08ld2cy 98 25 the the DT cord-346267-l08ld2cy 98 26 ratio ratio NN cord-346267-l08ld2cy 98 27 infinite infinite JJ cord-346267-l08ld2cy 98 28 . . . cord-346267-l08ld2cy 99 1 For for IN cord-346267-l08ld2cy 99 2 each each DT cord-346267-l08ld2cy 99 3 data data NN cord-346267-l08ld2cy 99 4 set set VBN cord-346267-l08ld2cy 99 5 , , , cord-346267-l08ld2cy 99 6 we -PRON- PRP cord-346267-l08ld2cy 99 7 obtained obtain VBD cord-346267-l08ld2cy 99 8 M M NNP cord-346267-l08ld2cy 99 9 = = SYM cord-346267-l08ld2cy 99 10 1 1 CD cord-346267-l08ld2cy 99 11 , , , cord-346267-l08ld2cy 99 12 000 000 CD cord-346267-l08ld2cy 99 13 samples sample NNS cord-346267-l08ld2cy 99 14 from from IN cord-346267-l08ld2cy 99 15 the the DT cord-346267-l08ld2cy 99 16 approximate approximate JJ cord-346267-l08ld2cy 99 17 joint joint JJ cord-346267-l08ld2cy 99 18 distribution distribution NN cord-346267-l08ld2cy 99 19 of of IN cord-346267-l08ld2cy 99 20 model model NN cord-346267-l08ld2cy 99 21 parameter parameter NN cord-346267-l08ld2cy 99 22 estimates estimate NNS cord-346267-l08ld2cy 99 23 using use VBG cord-346267-l08ld2cy 99 24 a a DT cord-346267-l08ld2cy 99 25 modified modify VBN cord-346267-l08ld2cy 99 26 Latin Latin NNP cord-346267-l08ld2cy 99 27 hypercube hypercube NN cord-346267-l08ld2cy 99 28 sampling sampling NN cord-346267-l08ld2cy 99 29 importance importance NN cord-346267-l08ld2cy 99 30 resampling resample VBG cord-346267-l08ld2cy 99 31 scheme scheme NN cord-346267-l08ld2cy 99 32 , , , cord-346267-l08ld2cy 99 33 described describe VBN cord-346267-l08ld2cy 99 34 in in IN cord-346267-l08ld2cy 99 35 detail detail NN cord-346267-l08ld2cy 99 36 elsewhere elsewhere RB cord-346267-l08ld2cy 99 37 ( ( -LRB- cord-346267-l08ld2cy 99 38 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 99 39 et et FW cord-346267-l08ld2cy 99 40 al al NNP cord-346267-l08ld2cy 99 41 . . NNP cord-346267-l08ld2cy 99 42 2010 2010 CD cord-346267-l08ld2cy 99 43 ) ) -RRB- cord-346267-l08ld2cy 99 44 . . . cord-346267-l08ld2cy 100 1 The the DT cord-346267-l08ld2cy 100 2 approach approach NN cord-346267-l08ld2cy 100 3 is be VBZ cord-346267-l08ld2cy 100 4 meant mean VBN cord-346267-l08ld2cy 100 5 to to TO cord-346267-l08ld2cy 100 6 quickly quickly RB cord-346267-l08ld2cy 100 7 obtain obtain VB cord-346267-l08ld2cy 100 8 an an DT cord-346267-l08ld2cy 100 9 approximate approximate JJ cord-346267-l08ld2cy 100 10 joint joint JJ cord-346267-l08ld2cy 100 11 distribution distribution NN cord-346267-l08ld2cy 100 12 of of IN cord-346267-l08ld2cy 100 13 maximum maximum JJ cord-346267-l08ld2cy 100 14 likelihood likelihood NN cord-346267-l08ld2cy 100 15 parameter parameter NN cord-346267-l08ld2cy 100 16 estimators estimator NNS cord-346267-l08ld2cy 100 17 . . . cord-346267-l08ld2cy 101 1 First first RB cord-346267-l08ld2cy 101 2 , , , cord-346267-l08ld2cy 101 3 an an DT cord-346267-l08ld2cy 101 4 area area NN cord-346267-l08ld2cy 101 5 of of IN cord-346267-l08ld2cy 101 6 parameter parameter NN cord-346267-l08ld2cy 101 7 space space NN cord-346267-l08ld2cy 101 8 to to TO cord-346267-l08ld2cy 101 9 be be VB cord-346267-l08ld2cy 101 10 sampled sample VBN cord-346267-l08ld2cy 101 11 is be VBZ cord-346267-l08ld2cy 101 12 defined define VBN cord-346267-l08ld2cy 101 13 by by IN cord-346267-l08ld2cy 101 14 constructing construct VBG cord-346267-l08ld2cy 101 15 a a DT cord-346267-l08ld2cy 101 16 d d NN cord-346267-l08ld2cy 101 17 -dimensional -dimensional JJ cord-346267-l08ld2cy 101 18 rectangle rectangle NN cord-346267-l08ld2cy 101 19 , , , cord-346267-l08ld2cy 101 20 in in IN cord-346267-l08ld2cy 101 21 which which WDT cord-346267-l08ld2cy 101 22 each each DT cord-346267-l08ld2cy 101 23 dimension dimension NN cord-346267-l08ld2cy 101 24 represents represent VBZ cord-346267-l08ld2cy 101 25 a a DT cord-346267-l08ld2cy 101 26 single single JJ cord-346267-l08ld2cy 101 27 model model NN cord-346267-l08ld2cy 101 28 parameter parameter NN cord-346267-l08ld2cy 101 29 , , , cord-346267-l08ld2cy 101 30 the the DT cord-346267-l08ld2cy 101 31 corresponding correspond VBG cord-346267-l08ld2cy 101 32 coordinate coordinate JJ cord-346267-l08ld2cy 101 33 interval interval NN cord-346267-l08ld2cy 101 34 is be VBZ cord-346267-l08ld2cy 101 35 centered center VBN cord-346267-l08ld2cy 101 36 on on IN cord-346267-l08ld2cy 101 37 the the DT cord-346267-l08ld2cy 101 38 maximum maximum JJ cord-346267-l08ld2cy 101 39 likelihood likelihood NN cord-346267-l08ld2cy 101 40 estimate estimate NN cord-346267-l08ld2cy 101 41 of of IN cord-346267-l08ld2cy 101 42 the the DT cord-346267-l08ld2cy 101 43 parameter parameter NN cord-346267-l08ld2cy 101 44 , , , cord-346267-l08ld2cy 101 45 and and CC cord-346267-l08ld2cy 101 46 the the DT cord-346267-l08ld2cy 101 47 lower low JJR cord-346267-l08ld2cy 101 48 and and CC cord-346267-l08ld2cy 101 49 upper upper JJ cord-346267-l08ld2cy 101 50 bounds bound NNS cord-346267-l08ld2cy 101 51 are be VBP cord-346267-l08ld2cy 101 52 determined determine VBN cord-346267-l08ld2cy 101 53 by by IN cord-346267-l08ld2cy 101 54 profile profile JJ cord-346267-l08ld2cy 101 55 likelihood likelihood NN cord-346267-l08ld2cy 101 56 . . . cord-346267-l08ld2cy 102 1 Second second RB cord-346267-l08ld2cy 102 2 , , , cord-346267-l08ld2cy 102 3 each each DT cord-346267-l08ld2cy 102 4 coordinate coordinate JJ cord-346267-l08ld2cy 102 5 interval interval NN cord-346267-l08ld2cy 102 6 is be VBZ cord-346267-l08ld2cy 102 7 partitioned partition VBN cord-346267-l08ld2cy 102 8 into into IN cord-346267-l08ld2cy 102 9 N n NN cord-346267-l08ld2cy 102 10 = = SYM cord-346267-l08ld2cy 102 11 1 1 CD cord-346267-l08ld2cy 102 12 , , , cord-346267-l08ld2cy 102 13 000d 000d CD cord-346267-l08ld2cy 102 14 subintervals subinterval NNS cord-346267-l08ld2cy 102 15 of of IN cord-346267-l08ld2cy 102 16 approximately approximately RB cord-346267-l08ld2cy 102 17 equal equal JJ cord-346267-l08ld2cy 102 18 probability probability NN cord-346267-l08ld2cy 102 19 , , , cord-346267-l08ld2cy 102 20 based base VBN cord-346267-l08ld2cy 102 21 on on IN cord-346267-l08ld2cy 102 22 the the DT cord-346267-l08ld2cy 102 23 asymptotic asymptotic JJ cord-346267-l08ld2cy 102 24 normal normal JJ cord-346267-l08ld2cy 102 25 approximation approximation NN cord-346267-l08ld2cy 102 26 to to IN cord-346267-l08ld2cy 102 27 the the DT cord-346267-l08ld2cy 102 28 likelihood likelihood NN cord-346267-l08ld2cy 102 29 surface surface NN cord-346267-l08ld2cy 102 30 . . . cord-346267-l08ld2cy 103 1 Third third JJ cord-346267-l08ld2cy 103 2 , , , cord-346267-l08ld2cy 103 3 N n CD cord-346267-l08ld2cy 103 4 samples sample NNS cord-346267-l08ld2cy 103 5 are be VBP cord-346267-l08ld2cy 103 6 drawn draw VBN cord-346267-l08ld2cy 103 7 from from IN cord-346267-l08ld2cy 103 8 the the DT cord-346267-l08ld2cy 103 9 d d NNP cord-346267-l08ld2cy 103 10 -dimensional -dimensional JJ cord-346267-l08ld2cy 103 11 rectangle rectangle NN cord-346267-l08ld2cy 103 12 , , , cord-346267-l08ld2cy 103 13 using use VBG cord-346267-l08ld2cy 103 14 the the DT cord-346267-l08ld2cy 103 15 Latin Latin NNP cord-346267-l08ld2cy 103 16 Hypercube Hypercube NNP cord-346267-l08ld2cy 103 17 scheme scheme NN cord-346267-l08ld2cy 103 18 ( ( -LRB- cord-346267-l08ld2cy 103 19 i.e. i.e. FW cord-346267-l08ld2cy 103 20 , , , cord-346267-l08ld2cy 103 21 each each DT cord-346267-l08ld2cy 103 22 interval interval NN cord-346267-l08ld2cy 103 23 in in IN cord-346267-l08ld2cy 103 24 every every DT cord-346267-l08ld2cy 103 25 coordinate coordinate NN cord-346267-l08ld2cy 103 26 is be VBZ cord-346267-l08ld2cy 103 27 sampled sample VBN cord-346267-l08ld2cy 103 28 exactly exactly RB cord-346267-l08ld2cy 103 29 once once RB cord-346267-l08ld2cy 103 30 ) ) -RRB- cord-346267-l08ld2cy 103 31 . . . cord-346267-l08ld2cy 104 1 Fourth fourth RB cord-346267-l08ld2cy 104 2 , , , cord-346267-l08ld2cy 104 3 M M NNP cord-346267-l08ld2cy 104 4 N N NNP cord-346267-l08ld2cy 104 5 points point NNS cord-346267-l08ld2cy 104 6 are be VBP cord-346267-l08ld2cy 104 7 resampled resample VBN cord-346267-l08ld2cy 104 8 based base VBN cord-346267-l08ld2cy 104 9 on on IN cord-346267-l08ld2cy 104 10 their -PRON- PRP$ cord-346267-l08ld2cy 104 11 importance importance NN cord-346267-l08ld2cy 104 12 ( ( -LRB- cord-346267-l08ld2cy 104 13 normalized normalized JJ cord-346267-l08ld2cy 104 14 likelihood likelihood NN cord-346267-l08ld2cy 104 15 ) ) -RRB- cord-346267-l08ld2cy 104 16 , , , cord-346267-l08ld2cy 104 17 following follow VBG cord-346267-l08ld2cy 104 18 the the DT cord-346267-l08ld2cy 104 19 procedure procedure NN cord-346267-l08ld2cy 104 20 described describe VBN cord-346267-l08ld2cy 104 21 elsewhere elsewhere RB cord-346267-l08ld2cy 105 1 ( ( -LRB- cord-346267-l08ld2cy 105 2 Skare Skare NNP cord-346267-l08ld2cy 105 3 et et NNP cord-346267-l08ld2cy 105 4 al al NNP cord-346267-l08ld2cy 105 5 . . NNP cord-346267-l08ld2cy 105 6 2003 2003 CD cord-346267-l08ld2cy 105 7 ) ) -RRB- cord-346267-l08ld2cy 105 8 . . . cord-346267-l08ld2cy 106 1 Variance variance NN cord-346267-l08ld2cy 106 2 in in IN cord-346267-l08ld2cy 106 3 estimated estimate VBN cord-346267-l08ld2cy 106 4 branch branch NN cord-346267-l08ld2cy 106 5 lengths length NNS cord-346267-l08ld2cy 106 6 was be VBD cord-346267-l08ld2cy 106 7 computed compute VBN cord-346267-l08ld2cy 106 8 using use VBG cord-346267-l08ld2cy 106 9 this this DT cord-346267-l08ld2cy 106 10 sample sample NN cord-346267-l08ld2cy 106 11 . . . cord-346267-l08ld2cy 107 1 All all DT cord-346267-l08ld2cy 107 2 codon codon NN cord-346267-l08ld2cy 107 3 - - HYPH cord-346267-l08ld2cy 107 4 based base VBN cord-346267-l08ld2cy 107 5 analyses analysis NNS cord-346267-l08ld2cy 107 6 were be VBD cord-346267-l08ld2cy 107 7 performed perform VBN cord-346267-l08ld2cy 107 8 in in IN cord-346267-l08ld2cy 107 9 HyPhy HyPhy NNP cord-346267-l08ld2cy 107 10 v2.0020110301 v2.0020110301 CD cord-346267-l08ld2cy 107 11 ) ) -RRB- cord-346267-l08ld2cy 107 12 . . . cord-346267-l08ld2cy 108 1 We -PRON- PRP cord-346267-l08ld2cy 108 2 simulated simulate VBD cord-346267-l08ld2cy 108 3 codon codon NN cord-346267-l08ld2cy 108 4 sequences sequence NNS cord-346267-l08ld2cy 108 5 along along IN cord-346267-l08ld2cy 108 6 a a DT cord-346267-l08ld2cy 108 7 single single JJ cord-346267-l08ld2cy 108 8 branch branch NN cord-346267-l08ld2cy 108 9 using use VBG cord-346267-l08ld2cy 108 10 the the DT cord-346267-l08ld2cy 108 11 MG94 MG94 NNP cord-346267-l08ld2cy 108 12 × × NNP cord-346267-l08ld2cy 108 13 REV REV NNP cord-346267-l08ld2cy 108 14 codon codon NN cord-346267-l08ld2cy 108 15 substitution substitution NN cord-346267-l08ld2cy 108 16 model model NN cord-346267-l08ld2cy 108 17 ) ) -RRB- cord-346267-l08ld2cy 108 18 with with IN cord-346267-l08ld2cy 108 19 site site NN cord-346267-l08ld2cy 108 20 - - HYPH cord-346267-l08ld2cy 108 21 specific specific JJ cord-346267-l08ld2cy 108 22 β β NN cord-346267-l08ld2cy 108 23 and and CC cord-346267-l08ld2cy 108 24 α α NN cord-346267-l08ld2cy 108 25 values value NNS cord-346267-l08ld2cy 108 26 inferred infer VBN cord-346267-l08ld2cy 108 27 using use VBG cord-346267-l08ld2cy 108 28 a a DT cord-346267-l08ld2cy 108 29 fixed fix VBN cord-346267-l08ld2cy 108 30 effects effect NNS cord-346267-l08ld2cy 108 31 likelihood likelihood NN cord-346267-l08ld2cy 108 32 method method NN cord-346267-l08ld2cy 108 33 on on IN cord-346267-l08ld2cy 108 34 internal internal JJ cord-346267-l08ld2cy 108 35 branches branch NNS cord-346267-l08ld2cy 108 36 ( ( -LRB- cord-346267-l08ld2cy 108 37 IFEL IFEL NNP cord-346267-l08ld2cy 108 38 ) ) -RRB- cord-346267-l08ld2cy 109 1 ( ( -LRB- cord-346267-l08ld2cy 109 2 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 109 3 et et FW cord-346267-l08ld2cy 109 4 al al NNP cord-346267-l08ld2cy 109 5 . . . cord-346267-l08ld2cy 109 6 2006 2006 CD cord-346267-l08ld2cy 109 7 ) ) -RRB- cord-346267-l08ld2cy 109 8 from from IN cord-346267-l08ld2cy 109 9 each each DT cord-346267-l08ld2cy 109 10 of of IN cord-346267-l08ld2cy 109 11 the the DT cord-346267-l08ld2cy 109 12 three three CD cord-346267-l08ld2cy 109 13 viral viral JJ cord-346267-l08ld2cy 109 14 data data NN cord-346267-l08ld2cy 109 15 sets set NNS cord-346267-l08ld2cy 109 16 . . . cord-346267-l08ld2cy 110 1 In in IN cord-346267-l08ld2cy 110 2 this this DT cord-346267-l08ld2cy 110 3 method method NN cord-346267-l08ld2cy 110 4 , , , cord-346267-l08ld2cy 110 5 three three CD cord-346267-l08ld2cy 110 6 rates rate NNS cord-346267-l08ld2cy 110 7 are be VBP cord-346267-l08ld2cy 110 8 inferred infer VBN cord-346267-l08ld2cy 110 9 from from IN cord-346267-l08ld2cy 110 10 each each DT cord-346267-l08ld2cy 110 11 site site NN cord-346267-l08ld2cy 110 12 : : : cord-346267-l08ld2cy 111 1 α α NNP cord-346267-l08ld2cy 111 2 s s NNP cord-346267-l08ld2cy 111 3 , , , cord-346267-l08ld2cy 111 4 β β NN cord-346267-l08ld2cy 111 5 I I NNP cord-346267-l08ld2cy 111 6 s s NNP cord-346267-l08ld2cy 111 7 , , , cord-346267-l08ld2cy 111 8 and and CC cord-346267-l08ld2cy 111 9 β β NNP cord-346267-l08ld2cy 111 10 T T NNP cord-346267-l08ld2cy 111 11 s s NNP cord-346267-l08ld2cy 111 12 . . . cord-346267-l08ld2cy 112 1 Subscript Subscript NNP cord-346267-l08ld2cy 112 2 s s NNP cord-346267-l08ld2cy 112 3 indicates indicate VBZ cord-346267-l08ld2cy 112 4 the the DT cord-346267-l08ld2cy 112 5 explicit explicit JJ cord-346267-l08ld2cy 112 6 dependence dependence NN cord-346267-l08ld2cy 112 7 of of IN cord-346267-l08ld2cy 112 8 substitution substitution NN cord-346267-l08ld2cy 112 9 rates rate NNS cord-346267-l08ld2cy 112 10 on on IN cord-346267-l08ld2cy 112 11 the the DT cord-346267-l08ld2cy 112 12 site site NN cord-346267-l08ld2cy 112 13 , , , cord-346267-l08ld2cy 112 14 from from IN cord-346267-l08ld2cy 112 15 which which WDT cord-346267-l08ld2cy 112 16 they -PRON- PRP cord-346267-l08ld2cy 112 17 are be VBP cord-346267-l08ld2cy 112 18 being be VBG cord-346267-l08ld2cy 112 19 estimated estimate VBN cord-346267-l08ld2cy 112 20 . . . cord-346267-l08ld2cy 113 1 α α NNP cord-346267-l08ld2cy 113 2 s s NNP cord-346267-l08ld2cy 113 3 is be VBZ cord-346267-l08ld2cy 113 4 the the DT cord-346267-l08ld2cy 113 5 tree tree NN cord-346267-l08ld2cy 113 6 - - HYPH cord-346267-l08ld2cy 113 7 wide wide JJ cord-346267-l08ld2cy 113 8 synonymous synonymous JJ cord-346267-l08ld2cy 113 9 rate rate NN cord-346267-l08ld2cy 113 10 , , , cord-346267-l08ld2cy 113 11 β β NN cord-346267-l08ld2cy 113 12 I i NN cord-346267-l08ld2cy 113 13 s s NNP cord-346267-l08ld2cy 113 14 is be VBZ cord-346267-l08ld2cy 113 15 the the DT cord-346267-l08ld2cy 113 16 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 113 17 rate rate NN cord-346267-l08ld2cy 113 18 shared share VBN cord-346267-l08ld2cy 113 19 by by IN cord-346267-l08ld2cy 113 20 all all DT cord-346267-l08ld2cy 113 21 internal internal JJ cord-346267-l08ld2cy 113 22 branches branch NNS cord-346267-l08ld2cy 113 23 , , , cord-346267-l08ld2cy 113 24 and and CC cord-346267-l08ld2cy 113 25 β β NNP cord-346267-l08ld2cy 113 26 T T NNP cord-346267-l08ld2cy 113 27 s s NNP cord-346267-l08ld2cy 113 28 is be VBZ cord-346267-l08ld2cy 113 29 the the DT cord-346267-l08ld2cy 113 30 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 113 31 rate rate NN cord-346267-l08ld2cy 113 32 shared share VBN cord-346267-l08ld2cy 113 33 by by IN cord-346267-l08ld2cy 113 34 all all DT cord-346267-l08ld2cy 113 35 terminal terminal JJ cord-346267-l08ld2cy 113 36 branches branch NNS cord-346267-l08ld2cy 113 37 . . . cord-346267-l08ld2cy 114 1 Only only RB cord-346267-l08ld2cy 114 2 internal internal JJ cord-346267-l08ld2cy 114 3 branches branch NNS cord-346267-l08ld2cy 114 4 were be VBD cord-346267-l08ld2cy 114 5 used use VBN cord-346267-l08ld2cy 114 6 to to TO cord-346267-l08ld2cy 114 7 generate generate VB cord-346267-l08ld2cy 114 8 empirical empirical JJ cord-346267-l08ld2cy 114 9 selection selection NN cord-346267-l08ld2cy 114 10 profiles profile NNS cord-346267-l08ld2cy 114 11 because because IN cord-346267-l08ld2cy 114 12 substitutions substitution NNS cord-346267-l08ld2cy 114 13 along along IN cord-346267-l08ld2cy 114 14 tips tip NNS cord-346267-l08ld2cy 114 15 in in IN cord-346267-l08ld2cy 114 16 viral viral JJ cord-346267-l08ld2cy 114 17 phylogenies phylogeny NNS cord-346267-l08ld2cy 114 18 are be VBP cord-346267-l08ld2cy 114 19 frequently frequently RB cord-346267-l08ld2cy 114 20 deleterious deleterious JJ cord-346267-l08ld2cy 114 21 and and CC cord-346267-l08ld2cy 114 22 transient transient JJ cord-346267-l08ld2cy 114 23 ( ( -LRB- cord-346267-l08ld2cy 114 24 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 114 25 et et FW cord-346267-l08ld2cy 114 26 al al NNP cord-346267-l08ld2cy 114 27 . . . cord-346267-l08ld2cy 115 1 2006 2006 CD cord-346267-l08ld2cy 115 2 ; ; : cord-346267-l08ld2cy 115 3 Pybus Pybus NNP cord-346267-l08ld2cy 115 4 et et NNP cord-346267-l08ld2cy 115 5 al al NNP cord-346267-l08ld2cy 115 6 . . NNP cord-346267-l08ld2cy 115 7 2007 2007 CD cord-346267-l08ld2cy 115 8 ) ) -RRB- cord-346267-l08ld2cy 115 9 . . . cord-346267-l08ld2cy 116 1 Furthermore furthermore RB cord-346267-l08ld2cy 116 2 , , , cord-346267-l08ld2cy 116 3 we -PRON- PRP cord-346267-l08ld2cy 116 4 were be VBD cord-346267-l08ld2cy 116 5 primarily primarily RB cord-346267-l08ld2cy 116 6 interested interested JJ cord-346267-l08ld2cy 116 7 in in IN cord-346267-l08ld2cy 116 8 the the DT cord-346267-l08ld2cy 116 9 effects effect NNS cord-346267-l08ld2cy 116 10 of of IN cord-346267-l08ld2cy 116 11 selection selection NN cord-346267-l08ld2cy 116 12 on on IN cord-346267-l08ld2cy 116 13 long long JJ cord-346267-l08ld2cy 116 14 internal internal JJ cord-346267-l08ld2cy 116 15 branches branch NNS cord-346267-l08ld2cy 116 16 , , , cord-346267-l08ld2cy 116 17 and and CC cord-346267-l08ld2cy 116 18 the the DT cord-346267-l08ld2cy 116 19 IFEL IFEL NNP cord-346267-l08ld2cy 116 20 profile profile NN cord-346267-l08ld2cy 116 21 ( ( -LRB- cord-346267-l08ld2cy 117 1 α α NNP cord-346267-l08ld2cy 117 2 s s NNP cord-346267-l08ld2cy 117 3 and and CC cord-346267-l08ld2cy 117 4 β β NN cord-346267-l08ld2cy 118 1 I -PRON- PRP cord-346267-l08ld2cy 118 2 s s VBZ cord-346267-l08ld2cy 118 3 inferred infer VBD cord-346267-l08ld2cy 118 4 for for IN cord-346267-l08ld2cy 118 5 each each DT cord-346267-l08ld2cy 118 6 site site NN cord-346267-l08ld2cy 118 7 ) ) -RRB- cord-346267-l08ld2cy 118 8 was be VBD cord-346267-l08ld2cy 118 9 meant mean VBN cord-346267-l08ld2cy 118 10 to to TO cord-346267-l08ld2cy 118 11 recapitulate recapitulate VB cord-346267-l08ld2cy 118 12 the the DT cord-346267-l08ld2cy 118 13 evolutionary evolutionary JJ cord-346267-l08ld2cy 118 14 process process NN cord-346267-l08ld2cy 118 15 along along IN cord-346267-l08ld2cy 118 16 an an DT cord-346267-l08ld2cy 118 17 " " `` cord-346267-l08ld2cy 118 18 average average JJ cord-346267-l08ld2cy 118 19 " " '' cord-346267-l08ld2cy 118 20 internal internal JJ cord-346267-l08ld2cy 118 21 branch branch NN cord-346267-l08ld2cy 118 22 . . . cord-346267-l08ld2cy 119 1 Simulations simulation NNS cord-346267-l08ld2cy 119 2 were be VBD cord-346267-l08ld2cy 119 3 initialized initialize VBN cord-346267-l08ld2cy 119 4 using use VBG cord-346267-l08ld2cy 119 5 ancestral ancestral JJ cord-346267-l08ld2cy 119 6 nucleotide nucleotide JJ cord-346267-l08ld2cy 119 7 sequences sequence NNS cord-346267-l08ld2cy 119 8 inferred infer VBN cord-346267-l08ld2cy 119 9 with with IN cord-346267-l08ld2cy 119 10 marginal marginal JJ cord-346267-l08ld2cy 119 11 maximum maximum JJ cord-346267-l08ld2cy 119 12 likelihood likelihood NN cord-346267-l08ld2cy 119 13 reconstruction reconstruction NN cord-346267-l08ld2cy 119 14 ( ( -LRB- cord-346267-l08ld2cy 119 15 Yang Yang NNP cord-346267-l08ld2cy 119 16 et et NNP cord-346267-l08ld2cy 119 17 al al NNP cord-346267-l08ld2cy 119 18 . . NNP cord-346267-l08ld2cy 119 19 1995 1995 CD cord-346267-l08ld2cy 119 20 ) ) -RRB- cord-346267-l08ld2cy 119 21 in in IN cord-346267-l08ld2cy 119 22 HyPhy HyPhy NNP cord-346267-l08ld2cy 119 23 , , , cord-346267-l08ld2cy 119 24 using use VBG cord-346267-l08ld2cy 119 25 the the DT cord-346267-l08ld2cy 119 26 MG94 MG94 NNP cord-346267-l08ld2cy 119 27 × × NNP cord-346267-l08ld2cy 119 28 REV REV NNP cord-346267-l08ld2cy 119 29 codon codon NN cord-346267-l08ld2cy 119 30 substitution substitution NN cord-346267-l08ld2cy 119 31 model model NN cord-346267-l08ld2cy 119 32 on on IN cord-346267-l08ld2cy 119 33 the the DT cord-346267-l08ld2cy 119 34 MCC MCC NNP cord-346267-l08ld2cy 119 35 phylogeny phylogeny NN cord-346267-l08ld2cy 119 36 obtained obtain VBN cord-346267-l08ld2cy 119 37 from from IN cord-346267-l08ld2cy 119 38 BMCMC BMCMC NNP cord-346267-l08ld2cy 119 39 general general JJ cord-346267-l08ld2cy 119 40 time time NN cord-346267-l08ld2cy 119 41 reversible reversible JJ cord-346267-l08ld2cy 119 42 ( ( -LRB- cord-346267-l08ld2cy 119 43 GTR GTR NNP cord-346267-l08ld2cy 119 44 ) ) -RRB- cord-346267-l08ld2cy 119 45 + + CC cord-346267-l08ld2cy 119 46 Γ γ NN cord-346267-l08ld2cy 119 47 4 4 CD cord-346267-l08ld2cy 119 48 analyses analysis NNS cord-346267-l08ld2cy 119 49 . . . cord-346267-l08ld2cy 120 1 The the DT cord-346267-l08ld2cy 120 2 marginal marginal JJ cord-346267-l08ld2cy 120 3 ancestral ancestral JJ cord-346267-l08ld2cy 120 4 sequence sequence NN cord-346267-l08ld2cy 120 5 was be VBD cord-346267-l08ld2cy 120 6 used use VBN cord-346267-l08ld2cy 120 7 to to TO cord-346267-l08ld2cy 120 8 provide provide VB cord-346267-l08ld2cy 120 9 a a DT cord-346267-l08ld2cy 120 10 realistic realistic JJ cord-346267-l08ld2cy 120 11 starting starting NN cord-346267-l08ld2cy 120 12 point point NN cord-346267-l08ld2cy 120 13 for for IN cord-346267-l08ld2cy 120 14 our -PRON- PRP$ cord-346267-l08ld2cy 120 15 simulations simulation NNS cord-346267-l08ld2cy 120 16 . . . cord-346267-l08ld2cy 121 1 Ten ten CD cord-346267-l08ld2cy 121 2 thousand thousand CD cord-346267-l08ld2cy 121 3 replicates replicate NNS cord-346267-l08ld2cy 121 4 were be VBD cord-346267-l08ld2cy 121 5 generated generate VBN cord-346267-l08ld2cy 121 6 for for IN cord-346267-l08ld2cy 121 7 each each DT cord-346267-l08ld2cy 121 8 branch branch NN cord-346267-l08ld2cy 121 9 length length NN cord-346267-l08ld2cy 121 10 ranging range VBG cord-346267-l08ld2cy 121 11 from from IN cord-346267-l08ld2cy 121 12 0.01 0.01 CD cord-346267-l08ld2cy 121 13 to to IN cord-346267-l08ld2cy 121 14 100 100 CD cord-346267-l08ld2cy 121 15 expected expect VBN cord-346267-l08ld2cy 121 16 substitutions substitution NNS cord-346267-l08ld2cy 121 17 per per IN cord-346267-l08ld2cy 121 18 nucleotide nucleotide JJ cord-346267-l08ld2cy 121 19 site site NN cord-346267-l08ld2cy 121 20 and and CC cord-346267-l08ld2cy 121 21 analyzed analyze VBD cord-346267-l08ld2cy 121 22 under under IN cord-346267-l08ld2cy 121 23 the the DT cord-346267-l08ld2cy 121 24 GTR GTR NNP cord-346267-l08ld2cy 121 25 + + SYM cord-346267-l08ld2cy 121 26 Γ γ NN cord-346267-l08ld2cy 121 27 4 4 CD cord-346267-l08ld2cy 121 28 and and CC cord-346267-l08ld2cy 121 29 Dual dual JJ cord-346267-l08ld2cy 121 30 substitution substitution NN cord-346267-l08ld2cy 121 31 models model NNS cord-346267-l08ld2cy 121 32 . . . cord-346267-l08ld2cy 122 1 We -PRON- PRP cord-346267-l08ld2cy 122 2 chose choose VBD cord-346267-l08ld2cy 122 3 to to TO cord-346267-l08ld2cy 122 4 simulate simulate VB cord-346267-l08ld2cy 122 5 branches branch NNS cord-346267-l08ld2cy 122 6 based base VBN cord-346267-l08ld2cy 122 7 on on IN cord-346267-l08ld2cy 122 8 expected expect VBN cord-346267-l08ld2cy 122 9 substitutions substitution NNS cord-346267-l08ld2cy 122 10 per per IN cord-346267-l08ld2cy 122 11 nucleotide nucleotide JJ cord-346267-l08ld2cy 122 12 site site NN cord-346267-l08ld2cy 122 13 , , , cord-346267-l08ld2cy 122 14 instead instead RB cord-346267-l08ld2cy 122 15 of of IN cord-346267-l08ld2cy 122 16 per per IN cord-346267-l08ld2cy 122 17 unit unit NN cord-346267-l08ld2cy 122 18 time time NN cord-346267-l08ld2cy 122 19 , , , cord-346267-l08ld2cy 122 20 because because IN cord-346267-l08ld2cy 122 21 of of IN cord-346267-l08ld2cy 122 22 the the DT cord-346267-l08ld2cy 122 23 difficulty difficulty NN cord-346267-l08ld2cy 122 24 in in IN cord-346267-l08ld2cy 122 25 standardizing standardize VBG cord-346267-l08ld2cy 122 26 time time NN cord-346267-l08ld2cy 122 27 for for IN cord-346267-l08ld2cy 122 28 variable variable JJ cord-346267-l08ld2cy 122 29 rate rate NN cord-346267-l08ld2cy 122 30 parameters parameter NNS cord-346267-l08ld2cy 122 31 ( ( -LRB- cord-346267-l08ld2cy 122 32 i.e. i.e. FW cord-346267-l08ld2cy 122 33 , , , cord-346267-l08ld2cy 122 34 α α NNP cord-346267-l08ld2cy 122 35 s s NNP cord-346267-l08ld2cy 122 36 , , , cord-346267-l08ld2cy 122 37 β β NN cord-346267-l08ld2cy 122 38 I I NNP cord-346267-l08ld2cy 122 39 s s NNP cord-346267-l08ld2cy 122 40 , , , cord-346267-l08ld2cy 122 41 and and CC cord-346267-l08ld2cy 122 42 substitution substitution NN cord-346267-l08ld2cy 122 43 rate rate NN cord-346267-l08ld2cy 122 44 ) ) -RRB- cord-346267-l08ld2cy 122 45 in in IN cord-346267-l08ld2cy 122 46 a a DT cord-346267-l08ld2cy 122 47 reversible reversible JJ cord-346267-l08ld2cy 122 48 model model NN cord-346267-l08ld2cy 122 49 ; ; : cord-346267-l08ld2cy 122 50 the the DT cord-346267-l08ld2cy 122 51 expected expect VBN cord-346267-l08ld2cy 122 52 number number NN cord-346267-l08ld2cy 122 53 of of IN cord-346267-l08ld2cy 122 54 substitutions substitution NNS cord-346267-l08ld2cy 122 55 divided divide VBN cord-346267-l08ld2cy 122 56 by by IN cord-346267-l08ld2cy 122 57 the the DT cord-346267-l08ld2cy 122 58 substitution substitution NN cord-346267-l08ld2cy 122 59 rate rate NN cord-346267-l08ld2cy 122 60 can can MD cord-346267-l08ld2cy 122 61 be be VB cord-346267-l08ld2cy 122 62 a a DT cord-346267-l08ld2cy 122 63 proxy proxy NN cord-346267-l08ld2cy 122 64 for for IN cord-346267-l08ld2cy 122 65 time time NN cord-346267-l08ld2cy 122 66 . . . cord-346267-l08ld2cy 123 1 Using use VBG cord-346267-l08ld2cy 123 2 BMCMC BMCMC NNP cord-346267-l08ld2cy 123 3 analysis analysis NN cord-346267-l08ld2cy 123 4 , , , cord-346267-l08ld2cy 123 5 we -PRON- PRP cord-346267-l08ld2cy 123 6 inferred infer VBD cord-346267-l08ld2cy 123 7 the the DT cord-346267-l08ld2cy 123 8 substitution substitution NN cord-346267-l08ld2cy 123 9 rate rate NN cord-346267-l08ld2cy 123 10 and and CC cord-346267-l08ld2cy 123 11 root root NN cord-346267-l08ld2cy 123 12 age age NN cord-346267-l08ld2cy 123 13 under under IN cord-346267-l08ld2cy 123 14 a a DT cord-346267-l08ld2cy 123 15 standard standard JJ cord-346267-l08ld2cy 123 16 nucleotide nucleotide JJ cord-346267-l08ld2cy 123 17 substitution substitution NN cord-346267-l08ld2cy 123 18 model model NN cord-346267-l08ld2cy 123 19 ( ( -LRB- cord-346267-l08ld2cy 123 20 GTR GTR NNP cord-346267-l08ld2cy 123 21 + + CC cord-346267-l08ld2cy 123 22 Γ γ NN cord-346267-l08ld2cy 123 23 4 4 CD cord-346267-l08ld2cy 123 24 ) ) -RRB- cord-346267-l08ld2cy 123 25 for for IN cord-346267-l08ld2cy 123 26 each each DT cord-346267-l08ld2cy 123 27 of of IN cord-346267-l08ld2cy 123 28 our -PRON- PRP$ cord-346267-l08ld2cy 123 29 three three CD cord-346267-l08ld2cy 123 30 viral viral JJ cord-346267-l08ld2cy 123 31 data data NN cord-346267-l08ld2cy 123 32 sets set NNS cord-346267-l08ld2cy 123 33 : : : cord-346267-l08ld2cy 124 1 MeV MeV NNP cord-346267-l08ld2cy 124 2 / / SYM cord-346267-l08ld2cy 124 3 RPV RPV NNP cord-346267-l08ld2cy 124 4 / / SYM cord-346267-l08ld2cy 124 5 PPRV PPRV NNP cord-346267-l08ld2cy 124 6 nucleoprotein nucleoprotein NN cord-346267-l08ld2cy 124 7 , , , cord-346267-l08ld2cy 124 8 EBOV EBOV NNP cord-346267-l08ld2cy 124 9 glycoprotein glycoprotein NN cord-346267-l08ld2cy 124 10 , , , cord-346267-l08ld2cy 124 11 and and CC cord-346267-l08ld2cy 124 12 AIV AIV NNP cord-346267-l08ld2cy 124 13 neuraminidase neuraminidase NN cord-346267-l08ld2cy 124 14 ( ( -LRB- cord-346267-l08ld2cy 124 15 table table NN cord-346267-l08ld2cy 124 16 1 1 CD cord-346267-l08ld2cy 124 17 ) ) -RRB- cord-346267-l08ld2cy 124 18 . . . cord-346267-l08ld2cy 125 1 The the DT cord-346267-l08ld2cy 125 2 viral viral JJ cord-346267-l08ld2cy 125 3 lineages lineage NNS cord-346267-l08ld2cy 125 4 examined examine VBN cord-346267-l08ld2cy 125 5 here here RB cord-346267-l08ld2cy 125 6 had have VBD cord-346267-l08ld2cy 125 7 tMRCAs tmrca VBN cord-346267-l08ld2cy 125 8 ranging range VBG cord-346267-l08ld2cy 125 9 from from IN cord-346267-l08ld2cy 125 10 hundreds hundred NNS cord-346267-l08ld2cy 125 11 to to IN cord-346267-l08ld2cy 125 12 several several JJ cord-346267-l08ld2cy 125 13 thousand thousand CD cord-346267-l08ld2cy 125 14 years year NNS cord-346267-l08ld2cy 125 15 before before IN cord-346267-l08ld2cy 125 16 present present JJ cord-346267-l08ld2cy 125 17 . . . cord-346267-l08ld2cy 126 1 All all DT cord-346267-l08ld2cy 126 2 three three CD cord-346267-l08ld2cy 126 3 viruses virus NNS cord-346267-l08ld2cy 126 4 exhibited exhibit VBD cord-346267-l08ld2cy 126 5 rapid rapid JJ cord-346267-l08ld2cy 126 6 substitution substitution NN cord-346267-l08ld2cy 126 7 rates rate NNS cord-346267-l08ld2cy 126 8 , , , cord-346267-l08ld2cy 126 9 on on IN cord-346267-l08ld2cy 126 10 par par NN cord-346267-l08ld2cy 126 11 with with IN cord-346267-l08ld2cy 126 12 previous previous JJ cord-346267-l08ld2cy 126 13 estimates estimate NNS cord-346267-l08ld2cy 126 14 for for IN cord-346267-l08ld2cy 126 15 related relate VBN cord-346267-l08ld2cy 126 16 RNA RNA NNP cord-346267-l08ld2cy 126 17 viruses virus NNS cord-346267-l08ld2cy 126 18 ( ( -LRB- cord-346267-l08ld2cy 126 19 Duffy Duffy NNP cord-346267-l08ld2cy 126 20 et et NNP cord-346267-l08ld2cy 126 21 al al NNP cord-346267-l08ld2cy 126 22 . . NNP cord-346267-l08ld2cy 126 23 2008 2008 CD cord-346267-l08ld2cy 126 24 ) ) -RRB- cord-346267-l08ld2cy 126 25 . . . cord-346267-l08ld2cy 127 1 Our -PRON- PRP$ cord-346267-l08ld2cy 127 2 inferred inferred JJ cord-346267-l08ld2cy 127 3 substitution substitution NN cord-346267-l08ld2cy 127 4 rate rate NN cord-346267-l08ld2cy 127 5 for for IN cord-346267-l08ld2cy 127 6 the the DT cord-346267-l08ld2cy 127 7 MeV MeV NNP cord-346267-l08ld2cy 127 8 / / SYM cord-346267-l08ld2cy 127 9 RPV RPV NNP cord-346267-l08ld2cy 127 10 / / SYM cord-346267-l08ld2cy 127 11 PPRV PPRV NNP cord-346267-l08ld2cy 127 12 nucleoprotein nucleoprotein NN cord-346267-l08ld2cy 127 13 of of IN cord-346267-l08ld2cy 127 14 9.65 9.65 CD cord-346267-l08ld2cy 127 15 × × NN cord-346267-l08ld2cy 127 16 10 10 CD cord-346267-l08ld2cy 127 17 −4 −4 CD cord-346267-l08ld2cy 127 18 ( ( -LRB- cord-346267-l08ld2cy 127 19 6.00 6.00 CD cord-346267-l08ld2cy 127 20 × × NN cord-346267-l08ld2cy 127 21 10 10 CD cord-346267-l08ld2cy 127 22 −4 −4 CD cord-346267-l08ld2cy 127 23 -1.47 -1.47 CD cord-346267-l08ld2cy 127 24 × × NN cord-346267-l08ld2cy 127 25 10 10 CD cord-346267-l08ld2cy 127 26 −3 −3 NN cord-346267-l08ld2cy 127 27 ) ) -RRB- cord-346267-l08ld2cy 127 28 substitutions substitution NNS cord-346267-l08ld2cy 127 29 / / SYM cord-346267-l08ld2cy 127 30 site site NN cord-346267-l08ld2cy 127 31 / / SYM cord-346267-l08ld2cy 127 32 year year NN cord-346267-l08ld2cy 127 33 under under IN cord-346267-l08ld2cy 127 34 a a DT cord-346267-l08ld2cy 127 35 GTR GTR NNP cord-346267-l08ld2cy 127 36 + + SYM cord-346267-l08ld2cy 127 37 Γ γ NN cord-346267-l08ld2cy 127 38 4 4 CD cord-346267-l08ld2cy 127 39 model model NN cord-346267-l08ld2cy 127 40 was be VBD cord-346267-l08ld2cy 127 41 over over IN cord-346267-l08ld2cy 127 42 50 50 CD cord-346267-l08ld2cy 127 43 % % NN cord-346267-l08ld2cy 127 44 faster fast RBR cord-346267-l08ld2cy 127 45 than than IN cord-346267-l08ld2cy 127 46 the the DT cord-346267-l08ld2cy 127 47 rate rate NN cord-346267-l08ld2cy 127 48 recently recently RB cord-346267-l08ld2cy 127 49 reported report VBN cord-346267-l08ld2cy 127 50 by by IN cord-346267-l08ld2cy 127 51 Furuse Furuse NNP cord-346267-l08ld2cy 127 52 et et NNP cord-346267-l08ld2cy 127 53 al al NNP cord-346267-l08ld2cy 127 54 . . . cord-346267-l08ld2cy 128 1 ( ( -LRB- cord-346267-l08ld2cy 128 2 2010 2010 CD cord-346267-l08ld2cy 128 3 ) ) -RRB- cord-346267-l08ld2cy 128 4 : : : cord-346267-l08ld2cy 128 5 6.02 6.02 CD cord-346267-l08ld2cy 128 6 × × NN cord-346267-l08ld2cy 128 7 10 10 CD cord-346267-l08ld2cy 128 8 −4 −4 CD cord-346267-l08ld2cy 128 9 ( ( -LRB- cord-346267-l08ld2cy 128 10 3.62 3.62 CD cord-346267-l08ld2cy 128 11 × × NN cord-346267-l08ld2cy 128 12 10 10 CD cord-346267-l08ld2cy 128 13 −4 −4 CD cord-346267-l08ld2cy 128 14 -8.76 -8.76 CD cord-346267-l08ld2cy 128 15 × × NN cord-346267-l08ld2cy 128 16 10 10 CD cord-346267-l08ld2cy 128 17 −4 −4 CD cord-346267-l08ld2cy 128 18 ) ) -RRB- cord-346267-l08ld2cy 128 19 substitutions substitution NNS cord-346267-l08ld2cy 128 20 / / SYM cord-346267-l08ld2cy 128 21 site site NN cord-346267-l08ld2cy 128 22 / / SYM cord-346267-l08ld2cy 128 23 year year NN cord-346267-l08ld2cy 128 24 . . . cord-346267-l08ld2cy 129 1 Therefore therefore RB cord-346267-l08ld2cy 129 2 , , , cord-346267-l08ld2cy 129 3 we -PRON- PRP cord-346267-l08ld2cy 129 4 inferred infer VBD cord-346267-l08ld2cy 129 5 a a DT cord-346267-l08ld2cy 129 6 substantially substantially RB cord-346267-l08ld2cy 129 7 younger young JJR cord-346267-l08ld2cy 129 8 tMRCA tmrca NN cord-346267-l08ld2cy 129 9 for for IN cord-346267-l08ld2cy 129 10 MeV MeV NNP cord-346267-l08ld2cy 129 11 and and CC cord-346267-l08ld2cy 129 12 RPV RPV NNP cord-346267-l08ld2cy 129 13 : : : cord-346267-l08ld2cy 130 1 1490 1490 CD cord-346267-l08ld2cy 130 2 CE CE NNP cord-346267-l08ld2cy 130 3 ( ( -LRB- cord-346267-l08ld2cy 130 4 1130 1130 CD cord-346267-l08ld2cy 130 5 - - SYM cord-346267-l08ld2cy 130 6 1810 1810 CD cord-346267-l08ld2cy 130 7 CE CE NNP cord-346267-l08ld2cy 130 8 ) ) -RRB- cord-346267-l08ld2cy 130 9 , , , cord-346267-l08ld2cy 130 10 instead instead RB cord-346267-l08ld2cy 130 11 of of IN cord-346267-l08ld2cy 130 12 1171 1171 CD cord-346267-l08ld2cy 130 13 CE CE NNP cord-346267-l08ld2cy 130 14 ( ( -LRB- cord-346267-l08ld2cy 130 15 678 678 CD cord-346267-l08ld2cy 130 16 - - SYM cord-346267-l08ld2cy 130 17 1612 1612 CD cord-346267-l08ld2cy 130 18 CE ce NN cord-346267-l08ld2cy 130 19 ) ) -RRB- cord-346267-l08ld2cy 130 20 . . . cord-346267-l08ld2cy 131 1 We -PRON- PRP cord-346267-l08ld2cy 131 2 consider consider VBP cord-346267-l08ld2cy 131 3 our -PRON- PRP$ cord-346267-l08ld2cy 131 4 rate rate NN cord-346267-l08ld2cy 131 5 estimates estimate NNS cord-346267-l08ld2cy 131 6 more more RBR cord-346267-l08ld2cy 131 7 reliable reliable JJ cord-346267-l08ld2cy 131 8 for for IN cord-346267-l08ld2cy 131 9 three three CD cord-346267-l08ld2cy 131 10 reasons reason NNS cord-346267-l08ld2cy 131 11 . . . cord-346267-l08ld2cy 132 1 First first RB cord-346267-l08ld2cy 132 2 , , , cord-346267-l08ld2cy 132 3 our -PRON- PRP$ cord-346267-l08ld2cy 132 4 values value NNS cord-346267-l08ld2cy 132 5 are be VBP cord-346267-l08ld2cy 132 6 in in IN cord-346267-l08ld2cy 132 7 close close JJ cord-346267-l08ld2cy 132 8 agreement agreement NN cord-346267-l08ld2cy 132 9 with with IN cord-346267-l08ld2cy 132 10 previous previous JJ cord-346267-l08ld2cy 132 11 substitution substitution NN cord-346267-l08ld2cy 132 12 rate rate NN cord-346267-l08ld2cy 132 13 estimates estimate NNS cord-346267-l08ld2cy 132 14 in in IN cord-346267-l08ld2cy 132 15 MeV MeV NNP cord-346267-l08ld2cy 132 16 : : : cord-346267-l08ld2cy 132 17 8.69 8.69 CD cord-346267-l08ld2cy 132 18 × × NN cord-346267-l08ld2cy 132 19 10 10 CD cord-346267-l08ld2cy 132 20 −4 −4 CD cord-346267-l08ld2cy 132 21 ( ( -LRB- cord-346267-l08ld2cy 132 22 5.89 5.89 CD cord-346267-l08ld2cy 132 23 × × NN cord-346267-l08ld2cy 132 24 10 10 CD cord-346267-l08ld2cy 132 25 −4 −4 CD cord-346267-l08ld2cy 133 1 -1.13 -1.13 : cord-346267-l08ld2cy 134 1 × × NN cord-346267-l08ld2cy 134 2 10 10 CD cord-346267-l08ld2cy 134 3 −3 −3 NN cord-346267-l08ld2cy 134 4 ) ) -RRB- cord-346267-l08ld2cy 135 1 substitutions substitution NNS cord-346267-l08ld2cy 135 2 / / SYM cord-346267-l08ld2cy 135 3 site site NN cord-346267-l08ld2cy 135 4 / / SYM cord-346267-l08ld2cy 135 5 year year NN cord-346267-l08ld2cy 136 1 ( ( -LRB- cord-346267-l08ld2cy 136 2 Pomeroy Pomeroy NNP cord-346267-l08ld2cy 136 3 et et NNP cord-346267-l08ld2cy 136 4 al al NNP cord-346267-l08ld2cy 136 5 . . NNP cord-346267-l08ld2cy 136 6 2008 2008 CD cord-346267-l08ld2cy 136 7 ) ) -RRB- cord-346267-l08ld2cy 136 8 . . . cord-346267-l08ld2cy 137 1 Second second JJ cord-346267-l08ld2cy 137 2 , , , cord-346267-l08ld2cy 137 3 Furuse Furuse NNP cord-346267-l08ld2cy 137 4 et et NNP cord-346267-l08ld2cy 137 5 al al NNP cord-346267-l08ld2cy 137 6 . . . cord-346267-l08ld2cy 138 1 ( ( -LRB- cord-346267-l08ld2cy 138 2 2010 2010 CD cord-346267-l08ld2cy 138 3 ) ) -RRB- cord-346267-l08ld2cy 138 4 included include VBD cord-346267-l08ld2cy 138 5 both both DT cord-346267-l08ld2cy 138 6 MeV MeV NNP cord-346267-l08ld2cy 138 7 and and CC cord-346267-l08ld2cy 138 8 RPV RPV NNP cord-346267-l08ld2cy 138 9 vaccine vaccine NN cord-346267-l08ld2cy 138 10 strains strain NNS cord-346267-l08ld2cy 138 11 in in IN cord-346267-l08ld2cy 138 12 their -PRON- PRP$ cord-346267-l08ld2cy 138 13 BMCMC BMCMC NNP cord-346267-l08ld2cy 138 14 dating date VBG cord-346267-l08ld2cy 138 15 analysis analysis NN cord-346267-l08ld2cy 139 1 ( ( -LRB- cord-346267-l08ld2cy 139 2 Rota Rota NNP cord-346267-l08ld2cy 139 3 et et NNP cord-346267-l08ld2cy 139 4 al al NNP cord-346267-l08ld2cy 139 5 . . . cord-346267-l08ld2cy 139 6 1994 1994 CD cord-346267-l08ld2cy 139 7 ; ; : cord-346267-l08ld2cy 139 8 Baron Baron NNP cord-346267-l08ld2cy 139 9 et et NNP cord-346267-l08ld2cy 139 10 al al NNP cord-346267-l08ld2cy 139 11 . . . cord-346267-l08ld2cy 139 12 1996 1996 CD cord-346267-l08ld2cy 139 13 ) ) -RRB- cord-346267-l08ld2cy 139 14 , , , cord-346267-l08ld2cy 139 15 despite despite IN cord-346267-l08ld2cy 139 16 evidence evidence NN cord-346267-l08ld2cy 139 17 that that IN cord-346267-l08ld2cy 139 18 the the DT cord-346267-l08ld2cy 139 19 inclusion inclusion NN cord-346267-l08ld2cy 139 20 of of IN cord-346267-l08ld2cy 139 21 vaccine vaccine NN cord-346267-l08ld2cy 139 22 strains strain NNS cord-346267-l08ld2cy 139 23 can can MD cord-346267-l08ld2cy 139 24 lead lead VB cord-346267-l08ld2cy 139 25 to to IN cord-346267-l08ld2cy 139 26 bias bias NN cord-346267-l08ld2cy 139 27 in in IN cord-346267-l08ld2cy 139 28 rate rate NN cord-346267-l08ld2cy 139 29 estimation estimation NN cord-346267-l08ld2cy 139 30 ( ( -LRB- cord-346267-l08ld2cy 139 31 Bush Bush NNP cord-346267-l08ld2cy 139 32 et et NNP cord-346267-l08ld2cy 139 33 al al NNP cord-346267-l08ld2cy 139 34 . . . cord-346267-l08ld2cy 139 35 2000 2000 CD cord-346267-l08ld2cy 139 36 ; ; : cord-346267-l08ld2cy 139 37 Wertheim Wertheim NNP cord-346267-l08ld2cy 139 38 2010 2010 CD cord-346267-l08ld2cy 139 39 ) ) -RRB- cord-346267-l08ld2cy 139 40 . . . cord-346267-l08ld2cy 140 1 Third third RB cord-346267-l08ld2cy 140 2 , , , cord-346267-l08ld2cy 140 3 Furuse Furuse NNP cord-346267-l08ld2cy 140 4 et et NNP cord-346267-l08ld2cy 140 5 al al NNP cord-346267-l08ld2cy 140 6 . . . cord-346267-l08ld2cy 141 1 ( ( -LRB- cord-346267-l08ld2cy 141 2 2010 2010 CD cord-346267-l08ld2cy 141 3 ) ) -RRB- cord-346267-l08ld2cy 141 4 fitted fit VBD cord-346267-l08ld2cy 141 5 an an DT cord-346267-l08ld2cy 141 6 exponential exponential JJ cord-346267-l08ld2cy 141 7 growth growth NN cord-346267-l08ld2cy 141 8 coalescent coalescent NN cord-346267-l08ld2cy 141 9 prior prior RB cord-346267-l08ld2cy 141 10 to to IN cord-346267-l08ld2cy 141 11 viruses virus NNS cord-346267-l08ld2cy 141 12 whose whose WP$ cord-346267-l08ld2cy 141 13 effective effective JJ cord-346267-l08ld2cy 141 14 population population NN cord-346267-l08ld2cy 141 15 size size NN cord-346267-l08ld2cy 141 16 has have VBZ cord-346267-l08ld2cy 141 17 been be VBN cord-346267-l08ld2cy 141 18 declining decline VBG cord-346267-l08ld2cy 141 19 - - HYPH cord-346267-l08ld2cy 141 20 due due IN cord-346267-l08ld2cy 141 21 to to IN cord-346267-l08ld2cy 141 22 eradication eradication NN cord-346267-l08ld2cy 141 23 efforts effort NNS cord-346267-l08ld2cy 141 24 against against IN cord-346267-l08ld2cy 141 25 both both DT cord-346267-l08ld2cy 141 26 MeV MeV NNP cord-346267-l08ld2cy 141 27 and and CC cord-346267-l08ld2cy 141 28 RPV RPV NNP cord-346267-l08ld2cy 141 29 ( ( -LRB- cord-346267-l08ld2cy 141 30 Moss Moss NNP cord-346267-l08ld2cy 141 31 2009 2009 CD cord-346267-l08ld2cy 141 32 ; ; : cord-346267-l08ld2cy 141 33 Normile Normile NNP cord-346267-l08ld2cy 141 34 2010 2010 CD cord-346267-l08ld2cy 141 35 ) ) -RRB- cord-346267-l08ld2cy 141 36 . . . cord-346267-l08ld2cy 142 1 Remarkably remarkably RB cord-346267-l08ld2cy 142 2 , , , cord-346267-l08ld2cy 142 3 our -PRON- PRP$ cord-346267-l08ld2cy 142 4 results result NNS cord-346267-l08ld2cy 142 5 are be VBP cord-346267-l08ld2cy 142 6 in in IN cord-346267-l08ld2cy 142 7 even even RB cord-346267-l08ld2cy 142 8 greater great JJR cord-346267-l08ld2cy 142 9 conflict conflict NN cord-346267-l08ld2cy 142 10 with with IN cord-346267-l08ld2cy 142 11 the the DT cord-346267-l08ld2cy 142 12 historical historical JJ cord-346267-l08ld2cy 142 13 documentation documentation NN cord-346267-l08ld2cy 142 14 of of IN cord-346267-l08ld2cy 142 15 measles measle NNS cord-346267-l08ld2cy 142 16 ( ( -LRB- cord-346267-l08ld2cy 142 17 Rāzī Rāzī NNP cord-346267-l08ld2cy 142 18 1848 1848 CD cord-346267-l08ld2cy 142 19 ) ) -RRB- cord-346267-l08ld2cy 142 20 , , , cord-346267-l08ld2cy 142 21 further further RB cord-346267-l08ld2cy 142 22 confirming confirm VBG cord-346267-l08ld2cy 142 23 our -PRON- PRP$ cord-346267-l08ld2cy 142 24 conjecture conjecture NN cord-346267-l08ld2cy 142 25 that that IN cord-346267-l08ld2cy 142 26 traditional traditional JJ cord-346267-l08ld2cy 142 27 nucleotide nucleotide JJ cord-346267-l08ld2cy 142 28 substitution substitution NN cord-346267-l08ld2cy 142 29 models model NNS cord-346267-l08ld2cy 142 30 can can MD cord-346267-l08ld2cy 142 31 underestimate underestimate VB cord-346267-l08ld2cy 142 32 the the DT cord-346267-l08ld2cy 142 33 age age NN cord-346267-l08ld2cy 142 34 of of IN cord-346267-l08ld2cy 142 35 ancient ancient JJ cord-346267-l08ld2cy 142 36 viral viral JJ cord-346267-l08ld2cy 142 37 divergence divergence NN cord-346267-l08ld2cy 142 38 events event NNS cord-346267-l08ld2cy 142 39 . . . cord-346267-l08ld2cy 143 1 We -PRON- PRP cord-346267-l08ld2cy 143 2 then then RB cord-346267-l08ld2cy 143 3 explored explore VBD cord-346267-l08ld2cy 143 4 whether whether IN cord-346267-l08ld2cy 143 5 alternative alternative JJ cord-346267-l08ld2cy 143 6 evolutionary evolutionary JJ cord-346267-l08ld2cy 143 7 models model NNS cord-346267-l08ld2cy 143 8 that that WDT cord-346267-l08ld2cy 143 9 , , , cord-346267-l08ld2cy 143 10 to to IN cord-346267-l08ld2cy 143 11 varying vary VBG cord-346267-l08ld2cy 143 12 extents extent NNS cord-346267-l08ld2cy 143 13 , , , cord-346267-l08ld2cy 143 14 account account VBP cord-346267-l08ld2cy 143 15 for for IN cord-346267-l08ld2cy 143 16 codon codon NN cord-346267-l08ld2cy 143 17 structure structure NN cord-346267-l08ld2cy 143 18 were be VBD cord-346267-l08ld2cy 143 19 able able JJ cord-346267-l08ld2cy 143 20 to to TO cord-346267-l08ld2cy 143 21 recover recover VB cord-346267-l08ld2cy 143 22 the the DT cord-346267-l08ld2cy 143 23 expected expect VBN cord-346267-l08ld2cy 143 24 older old JJR cord-346267-l08ld2cy 143 25 tMRCAs tmrca VBD cord-346267-l08ld2cy 143 26 for for IN cord-346267-l08ld2cy 143 27 these these DT cord-346267-l08ld2cy 143 28 viruses virus NNS cord-346267-l08ld2cy 143 29 ( ( -LRB- cord-346267-l08ld2cy 143 30 table table NN cord-346267-l08ld2cy 143 31 1 1 CD cord-346267-l08ld2cy 143 32 ) ) -RRB- cord-346267-l08ld2cy 143 33 . . . cord-346267-l08ld2cy 144 1 The the DT cord-346267-l08ld2cy 144 2 simplest simple JJS cord-346267-l08ld2cy 144 3 approach approach NN cord-346267-l08ld2cy 144 4 , , , cord-346267-l08ld2cy 144 5 excluding exclude VBG cord-346267-l08ld2cy 144 6 the the DT cord-346267-l08ld2cy 144 7 third third JJ cord-346267-l08ld2cy 144 8 codon codon NN cord-346267-l08ld2cy 144 9 position position NN cord-346267-l08ld2cy 144 10 in in IN cord-346267-l08ld2cy 144 11 a a DT cord-346267-l08ld2cy 144 12 GTR GTR NNP cord-346267-l08ld2cy 144 13 + + SYM cord-346267-l08ld2cy 144 14 Γ γ NN cord-346267-l08ld2cy 144 15 4 4 CD cord-346267-l08ld2cy 144 16 model model NN cord-346267-l08ld2cy 144 17 , , , cord-346267-l08ld2cy 144 18 has have VBZ cord-346267-l08ld2cy 144 19 been be VBN cord-346267-l08ld2cy 144 20 used use VBN cord-346267-l08ld2cy 144 21 to to TO cord-346267-l08ld2cy 144 22 remove remove VB cord-346267-l08ld2cy 144 23 synonymous synonymous JJ cord-346267-l08ld2cy 144 24 sites site NNS cord-346267-l08ld2cy 144 25 that that WDT cord-346267-l08ld2cy 144 26 might may MD cord-346267-l08ld2cy 144 27 have have VB cord-346267-l08ld2cy 144 28 experienced experience VBN cord-346267-l08ld2cy 144 29 saturation saturation NN cord-346267-l08ld2cy 144 30 ( ( -LRB- cord-346267-l08ld2cy 144 31 Worobey Worobey NNP cord-346267-l08ld2cy 144 32 et et NNP cord-346267-l08ld2cy 144 33 al al NNP cord-346267-l08ld2cy 144 34 . . NNP cord-346267-l08ld2cy 144 35 2010 2010 CD cord-346267-l08ld2cy 144 36 ) ) -RRB- cord-346267-l08ld2cy 144 37 . . . cord-346267-l08ld2cy 145 1 A a DT cord-346267-l08ld2cy 145 2 more more RBR cord-346267-l08ld2cy 145 3 sophisticated sophisticated JJ cord-346267-l08ld2cy 145 4 approach approach NN cord-346267-l08ld2cy 145 5 , , , cord-346267-l08ld2cy 145 6 the the DT cord-346267-l08ld2cy 145 7 SRD06 SRD06 NNP cord-346267-l08ld2cy 145 8 model model NN cord-346267-l08ld2cy 145 9 ( ( -LRB- cord-346267-l08ld2cy 145 10 Shapiro Shapiro NNP cord-346267-l08ld2cy 145 11 et et NNP cord-346267-l08ld2cy 145 12 al al NNP cord-346267-l08ld2cy 145 13 . . . cord-346267-l08ld2cy 145 14 2006 2006 CD cord-346267-l08ld2cy 145 15 ) ) -RRB- cord-346267-l08ld2cy 145 16 , , , cord-346267-l08ld2cy 145 17 allows allow VBZ cord-346267-l08ld2cy 145 18 first first JJ cord-346267-l08ld2cy 145 19 and and CC cord-346267-l08ld2cy 145 20 second second JJ cord-346267-l08ld2cy 145 21 codon codon NN cord-346267-l08ld2cy 145 22 positions position NNS cord-346267-l08ld2cy 145 23 to to TO cord-346267-l08ld2cy 145 24 have have VB cord-346267-l08ld2cy 145 25 a a DT cord-346267-l08ld2cy 145 26 different different JJ cord-346267-l08ld2cy 145 27 transition transition NN cord-346267-l08ld2cy 145 28 / / SYM cord-346267-l08ld2cy 145 29 transversion transversion NN cord-346267-l08ld2cy 145 30 ratio ratio NN cord-346267-l08ld2cy 145 31 and and CC cord-346267-l08ld2cy 145 32 Γ γ NN cord-346267-l08ld2cy 145 33 4 4 CD cord-346267-l08ld2cy 145 34 shape shape NN cord-346267-l08ld2cy 145 35 parameter parameter NN cord-346267-l08ld2cy 145 36 from from IN cord-346267-l08ld2cy 145 37 the the DT cord-346267-l08ld2cy 145 38 third third JJ cord-346267-l08ld2cy 145 39 position position NN cord-346267-l08ld2cy 145 40 . . . cord-346267-l08ld2cy 146 1 Amino amino NN cord-346267-l08ld2cy 146 2 acid acid NN cord-346267-l08ld2cy 146 3 substitution substitution NN cord-346267-l08ld2cy 146 4 models model NNS cord-346267-l08ld2cy 146 5 have have VBP cord-346267-l08ld2cy 146 6 also also RB cord-346267-l08ld2cy 146 7 been be VBN cord-346267-l08ld2cy 146 8 successfully successfully RB cord-346267-l08ld2cy 146 9 implemented implement VBN cord-346267-l08ld2cy 146 10 to to TO cord-346267-l08ld2cy 146 11 estimate estimate VB cord-346267-l08ld2cy 146 12 the the DT cord-346267-l08ld2cy 146 13 age age NN cord-346267-l08ld2cy 146 14 of of IN cord-346267-l08ld2cy 146 15 RNA RNA NNP cord-346267-l08ld2cy 146 16 viruses virus NNS cord-346267-l08ld2cy 146 17 ( ( -LRB- cord-346267-l08ld2cy 146 18 Zlateva Zlateva NNP cord-346267-l08ld2cy 146 19 et et FW cord-346267-l08ld2cy 146 20 al al NNP cord-346267-l08ld2cy 146 21 . . NNP cord-346267-l08ld2cy 146 22 2005 2005 CD cord-346267-l08ld2cy 146 23 ; ; : cord-346267-l08ld2cy 146 24 Wertheim Wertheim NNP cord-346267-l08ld2cy 146 25 et et NNP cord-346267-l08ld2cy 146 26 al al NNP cord-346267-l08ld2cy 146 27 . . NNP cord-346267-l08ld2cy 146 28 2009 2009 CD cord-346267-l08ld2cy 146 29 ) ) -RRB- cord-346267-l08ld2cy 146 30 . . . cord-346267-l08ld2cy 147 1 Finally finally RB cord-346267-l08ld2cy 147 2 , , , cord-346267-l08ld2cy 147 3 we -PRON- PRP cord-346267-l08ld2cy 147 4 evaluated evaluate VBD cord-346267-l08ld2cy 147 5 an an DT cord-346267-l08ld2cy 147 6 available available JJ cord-346267-l08ld2cy 147 7 codon codon NN cord-346267-l08ld2cy 147 8 - - HYPH cord-346267-l08ld2cy 147 9 based base VBN cord-346267-l08ld2cy 147 10 substitution substitution NN cord-346267-l08ld2cy 147 11 model model NN cord-346267-l08ld2cy 147 12 , , , cord-346267-l08ld2cy 147 13 GY94 GY94 NNP cord-346267-l08ld2cy 147 14 ( ( -LRB- cord-346267-l08ld2cy 147 15 Goldman Goldman NNP cord-346267-l08ld2cy 147 16 and and CC cord-346267-l08ld2cy 147 17 Yang Yang NNP cord-346267-l08ld2cy 147 18 1994 1994 CD cord-346267-l08ld2cy 147 19 ) ) -RRB- cord-346267-l08ld2cy 147 20 . . . cord-346267-l08ld2cy 148 1 Generally generally RB cord-346267-l08ld2cy 148 2 , , , cord-346267-l08ld2cy 148 3 these these DT cord-346267-l08ld2cy 148 4 alternate alternate JJ cord-346267-l08ld2cy 148 5 models model NNS cord-346267-l08ld2cy 148 6 either either CC cord-346267-l08ld2cy 148 7 produced produce VBD cord-346267-l08ld2cy 148 8 younger young JJR cord-346267-l08ld2cy 148 9 tMRCAs tmrcas IN cord-346267-l08ld2cy 148 10 ( ( -LRB- cord-346267-l08ld2cy 148 11 e.g. e.g. RB cord-346267-l08ld2cy 148 12 , , , cord-346267-l08ld2cy 148 13 GTR GTR NNP cord-346267-l08ld2cy 148 14 + + CC cord-346267-l08ld2cy 148 15 Γ γ NN cord-346267-l08ld2cy 148 16 4 4 CD cord-346267-l08ld2cy 148 17 excluding exclude VBG cord-346267-l08ld2cy 148 18 third third JJ cord-346267-l08ld2cy 148 19 positions position NNS cord-346267-l08ld2cy 148 20 and and CC cord-346267-l08ld2cy 148 21 Whelan Whelan NNP cord-346267-l08ld2cy 148 22 and and CC cord-346267-l08ld2cy 148 23 Goldman Goldman NNP cord-346267-l08ld2cy 148 24 + + CC cord-346267-l08ld2cy 148 25 Γ Γ NNP cord-346267-l08ld2cy 148 26 4 4 CD cord-346267-l08ld2cy 148 27 ) ) -RRB- cord-346267-l08ld2cy 148 28 or or CC cord-346267-l08ld2cy 148 29 tMRCAs tmrca VBN cord-346267-l08ld2cy 148 30 that that WDT cord-346267-l08ld2cy 148 31 were be VBD cord-346267-l08ld2cy 148 32 indistinguishable indistinguishable JJ cord-346267-l08ld2cy 148 33 from from IN cord-346267-l08ld2cy 148 34 GTR GTR NNP cord-346267-l08ld2cy 148 35 + + CC cord-346267-l08ld2cy 148 36 Γ γ NN cord-346267-l08ld2cy 148 37 4 4 CD cord-346267-l08ld2cy 148 38 ( ( -LRB- cord-346267-l08ld2cy 148 39 e.g. e.g. RB cord-346267-l08ld2cy 148 40 , , , cord-346267-l08ld2cy 148 41 SRD06 SRD06 NNP cord-346267-l08ld2cy 148 42 and and CC cord-346267-l08ld2cy 148 43 GY94 GY94 NNP cord-346267-l08ld2cy 148 44 + + CC cord-346267-l08ld2cy 148 45 Γ Γ NNP cord-346267-l08ld2cy 148 46 4 4 CD cord-346267-l08ld2cy 148 47 ) ) -RRB- cord-346267-l08ld2cy 148 48 . . . cord-346267-l08ld2cy 149 1 A a DT cord-346267-l08ld2cy 149 2 possible possible JJ cord-346267-l08ld2cy 149 3 exception exception NN cord-346267-l08ld2cy 149 4 was be VBD cord-346267-l08ld2cy 149 5 analysis analysis NN cord-346267-l08ld2cy 149 6 of of IN cord-346267-l08ld2cy 149 7 the the DT cord-346267-l08ld2cy 149 8 EBOV EBOV NNP cord-346267-l08ld2cy 149 9 data datum NNS cord-346267-l08ld2cy 149 10 set set VBN cord-346267-l08ld2cy 149 11 with with IN cord-346267-l08ld2cy 149 12 GY94 GY94 NNP cord-346267-l08ld2cy 149 13 + + CC cord-346267-l08ld2cy 149 14 Γ γ NN cord-346267-l08ld2cy 149 15 4 4 CD cord-346267-l08ld2cy 149 16 , , , cord-346267-l08ld2cy 149 17 which which WDT cord-346267-l08ld2cy 149 18 produced produce VBD cord-346267-l08ld2cy 149 19 a a DT cord-346267-l08ld2cy 149 20 root root NN cord-346267-l08ld2cy 149 21 tMRCA tmrca NN cord-346267-l08ld2cy 149 22 that that WDT cord-346267-l08ld2cy 149 23 was be VBD cord-346267-l08ld2cy 149 24 50 50 CD cord-346267-l08ld2cy 149 25 % % NN cord-346267-l08ld2cy 149 26 older old JJR cord-346267-l08ld2cy 149 27 than than IN cord-346267-l08ld2cy 149 28 the the DT cord-346267-l08ld2cy 149 29 GTR GTR NNP cord-346267-l08ld2cy 149 30 + + SYM cord-346267-l08ld2cy 149 31 Γ γ NN cord-346267-l08ld2cy 149 32 4 4 CD cord-346267-l08ld2cy 149 33 estimate estimate NN cord-346267-l08ld2cy 149 34 . . . cord-346267-l08ld2cy 150 1 Finally finally RB cord-346267-l08ld2cy 150 2 , , , cord-346267-l08ld2cy 150 3 as as IN cord-346267-l08ld2cy 150 4 a a DT cord-346267-l08ld2cy 150 5 point point NN cord-346267-l08ld2cy 150 6 of of IN cord-346267-l08ld2cy 150 7 comparison comparison NN cord-346267-l08ld2cy 150 8 , , , cord-346267-l08ld2cy 150 9 we -PRON- PRP cord-346267-l08ld2cy 150 10 also also RB cord-346267-l08ld2cy 150 11 investigated investigate VBD cord-346267-l08ld2cy 150 12 how how WRB cord-346267-l08ld2cy 150 13 removing remove VBG cord-346267-l08ld2cy 150 14 site site NN cord-346267-l08ld2cy 150 15 - - HYPH cord-346267-l08ld2cy 150 16 to to IN cord-346267-l08ld2cy 150 17 - - HYPH cord-346267-l08ld2cy 150 18 site site NN cord-346267-l08ld2cy 150 19 rate rate NN cord-346267-l08ld2cy 150 20 variation variation NN cord-346267-l08ld2cy 150 21 affected affect VBN cord-346267-l08ld2cy 150 22 dating date VBG cord-346267-l08ld2cy 150 23 inference inference NN cord-346267-l08ld2cy 150 24 by by IN cord-346267-l08ld2cy 150 25 using use VBG cord-346267-l08ld2cy 150 26 a a DT cord-346267-l08ld2cy 150 27 GTR GTR NNP cord-346267-l08ld2cy 150 28 model model NN cord-346267-l08ld2cy 150 29 ( ( -LRB- cord-346267-l08ld2cy 150 30 Tavaré Tavaré NNP cord-346267-l08ld2cy 150 31 1986 1986 CD cord-346267-l08ld2cy 150 32 ) ) -RRB- cord-346267-l08ld2cy 150 33 ; ; : cord-346267-l08ld2cy 150 34 ignoring ignore VBG cord-346267-l08ld2cy 150 35 rate rate NN cord-346267-l08ld2cy 150 36 variation variation NN cord-346267-l08ld2cy 150 37 invariably invariably RB cord-346267-l08ld2cy 150 38 produced produce VBD cord-346267-l08ld2cy 150 39 younger young JJR cord-346267-l08ld2cy 150 40 tMRCAs tmrcas CD cord-346267-l08ld2cy 150 41 . . . cord-346267-l08ld2cy 151 1 We -PRON- PRP cord-346267-l08ld2cy 151 2 simulated simulate VBD cord-346267-l08ld2cy 151 3 codon codon NN cord-346267-l08ld2cy 151 4 sequences sequence NNS cord-346267-l08ld2cy 151 5 along along IN cord-346267-l08ld2cy 151 6 a a DT cord-346267-l08ld2cy 151 7 single single JJ cord-346267-l08ld2cy 151 8 branch branch NN cord-346267-l08ld2cy 151 9 under under IN cord-346267-l08ld2cy 151 10 a a DT cord-346267-l08ld2cy 151 11 substitution substitution NN cord-346267-l08ld2cy 151 12 model model NN cord-346267-l08ld2cy 151 13 with with IN cord-346267-l08ld2cy 151 14 site site NN cord-346267-l08ld2cy 151 15 - - HYPH cord-346267-l08ld2cy 151 16 to to IN cord-346267-l08ld2cy 151 17 - - HYPH cord-346267-l08ld2cy 151 18 site site NN cord-346267-l08ld2cy 151 19 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 151 20 rate rate NN cord-346267-l08ld2cy 151 21 variation variation NN cord-346267-l08ld2cy 151 22 and and CC cord-346267-l08ld2cy 151 23 inferred infer VBD cord-346267-l08ld2cy 151 24 the the DT cord-346267-l08ld2cy 151 25 length length NN cord-346267-l08ld2cy 151 26 of of IN cord-346267-l08ld2cy 151 27 the the DT cord-346267-l08ld2cy 151 28 branch branch NN cord-346267-l08ld2cy 151 29 using use VBG cord-346267-l08ld2cy 151 30 GTR GTR NNP cord-346267-l08ld2cy 151 31 + + CC cord-346267-l08ld2cy 151 32 Γ γ NN cord-346267-l08ld2cy 151 33 4 4 CD cord-346267-l08ld2cy 151 34 . . . cord-346267-l08ld2cy 152 1 An an DT cord-346267-l08ld2cy 152 2 increase increase NN cord-346267-l08ld2cy 152 3 in in IN cord-346267-l08ld2cy 152 4 the the DT cord-346267-l08ld2cy 152 5 proportion proportion NN cord-346267-l08ld2cy 152 6 of of IN cord-346267-l08ld2cy 152 7 sites site NNS cord-346267-l08ld2cy 152 8 simulated simulate VBN cord-346267-l08ld2cy 152 9 under under IN cord-346267-l08ld2cy 152 10 purifying purify VBG cord-346267-l08ld2cy 152 11 selection selection NN cord-346267-l08ld2cy 152 12 ( ( -LRB- cord-346267-l08ld2cy 152 13 i.e. i.e. FW cord-346267-l08ld2cy 152 14 , , , cord-346267-l08ld2cy 152 15 β β NN cord-346267-l08ld2cy 152 16 s s NN cord-346267-l08ld2cy 152 17 = = SYM cord-346267-l08ld2cy 152 18 0 0 CD cord-346267-l08ld2cy 152 19 ) ) -RRB- cord-346267-l08ld2cy 152 20 resulted result VBD cord-346267-l08ld2cy 152 21 in in IN cord-346267-l08ld2cy 152 22 shorter short JJR cord-346267-l08ld2cy 152 23 branches branch NNS cord-346267-l08ld2cy 152 24 inferred infer VBN cord-346267-l08ld2cy 152 25 by by IN cord-346267-l08ld2cy 152 26 a a DT cord-346267-l08ld2cy 152 27 GTR GTR NNP cord-346267-l08ld2cy 152 28 + + SYM cord-346267-l08ld2cy 152 29 Γ γ NN cord-346267-l08ld2cy 152 30 4 4 CD cord-346267-l08ld2cy 152 31 model model NN cord-346267-l08ld2cy 152 32 ( ( -LRB- cord-346267-l08ld2cy 152 33 fig fig NN cord-346267-l08ld2cy 152 34 . . . cord-346267-l08ld2cy 152 35 1A 1a LS cord-346267-l08ld2cy 152 36 ) ) -RRB- cord-346267-l08ld2cy 152 37 . . . cord-346267-l08ld2cy 153 1 Sufficiently sufficiently RB cord-346267-l08ld2cy 153 2 strong strong JJ cord-346267-l08ld2cy 153 3 purifying purifying NN cord-346267-l08ld2cy 153 4 selection selection NN cord-346267-l08ld2cy 153 5 could could MD cord-346267-l08ld2cy 153 6 theoretically theoretically RB cord-346267-l08ld2cy 153 7 lead lead VB cord-346267-l08ld2cy 153 8 to to IN cord-346267-l08ld2cy 153 9 underestimates underestimate NNS cord-346267-l08ld2cy 153 10 of of IN cord-346267-l08ld2cy 153 11 branch branch NN cord-346267-l08ld2cy 153 12 lengths length NNS cord-346267-l08ld2cy 153 13 by by IN cord-346267-l08ld2cy 153 14 an an DT cord-346267-l08ld2cy 153 15 order order NN cord-346267-l08ld2cy 153 16 of of IN cord-346267-l08ld2cy 153 17 magnitude magnitude NN cord-346267-l08ld2cy 153 18 because because IN cord-346267-l08ld2cy 153 19 purifying purify VBG cord-346267-l08ld2cy 153 20 selection selection NN cord-346267-l08ld2cy 153 21 slows slow VBZ cord-346267-l08ld2cy 153 22 down down IN cord-346267-l08ld2cy 153 23 the the DT cord-346267-l08ld2cy 153 24 rate rate NN cord-346267-l08ld2cy 153 25 of of IN cord-346267-l08ld2cy 153 26 evolution evolution NN cord-346267-l08ld2cy 153 27 relative relative JJ cord-346267-l08ld2cy 153 28 to to IN cord-346267-l08ld2cy 153 29 the the DT cord-346267-l08ld2cy 153 30 synonymous synonymous JJ cord-346267-l08ld2cy 153 31 rate rate NN cord-346267-l08ld2cy 153 32 . . . cord-346267-l08ld2cy 154 1 At at IN cord-346267-l08ld2cy 154 2 the the DT cord-346267-l08ld2cy 154 3 extreme extreme NN cord-346267-l08ld2cy 154 4 ( ( -LRB- cord-346267-l08ld2cy 154 5 β β NN cord-346267-l08ld2cy 154 6 s s NN cord-346267-l08ld2cy 154 7 = = SYM cord-346267-l08ld2cy 154 8 0 0 CD cord-346267-l08ld2cy 154 9 , , , cord-346267-l08ld2cy 154 10 branch branch NN cord-346267-l08ld2cy 154 11 length length NN cord-346267-l08ld2cy 154 12 → → IN cord-346267-l08ld2cy 154 13 ∞ ∞ NNP cord-346267-l08ld2cy 154 14 ) ) -RRB- cord-346267-l08ld2cy 154 15 , , , cord-346267-l08ld2cy 154 16 one one NN cord-346267-l08ld2cy 154 17 obtains obtain VBZ cord-346267-l08ld2cy 154 18 perfectly perfectly RB cord-346267-l08ld2cy 154 19 conserved conserve VBN cord-346267-l08ld2cy 154 20 amino amino NN cord-346267-l08ld2cy 154 21 acid acid NN cord-346267-l08ld2cy 154 22 sequences sequence NNS cord-346267-l08ld2cy 154 23 and and CC cord-346267-l08ld2cy 154 24 completely completely RB cord-346267-l08ld2cy 154 25 saturated saturated JJ cord-346267-l08ld2cy 154 26 synonymous synonymous JJ cord-346267-l08ld2cy 154 27 substitutions substitution NNS cord-346267-l08ld2cy 154 28 . . . cord-346267-l08ld2cy 155 1 For for IN cord-346267-l08ld2cy 155 2 sequences sequence NNS cord-346267-l08ld2cy 155 3 with with IN cord-346267-l08ld2cy 155 4 10 10 CD cord-346267-l08ld2cy 155 5 % % NN cord-346267-l08ld2cy 155 6 of of IN cord-346267-l08ld2cy 155 7 sites site NNS cord-346267-l08ld2cy 155 8 under under IN cord-346267-l08ld2cy 155 9 strict strict JJ cord-346267-l08ld2cy 155 10 conservation conservation NN cord-346267-l08ld2cy 155 11 , , , cord-346267-l08ld2cy 155 12 the the DT cord-346267-l08ld2cy 155 13 nucleotide nucleotide JJ cord-346267-l08ld2cy 155 14 estimator estimator NN cord-346267-l08ld2cy 155 15 approached approach VBD cord-346267-l08ld2cy 155 16 the the DT cord-346267-l08ld2cy 155 17 asymptote asymptote NN cord-346267-l08ld2cy 155 18 of of IN cord-346267-l08ld2cy 155 19 7.1 7.1 CD cord-346267-l08ld2cy 155 20 substitutions substitution NNS cord-346267-l08ld2cy 155 21 / / SYM cord-346267-l08ld2cy 155 22 site site NN cord-346267-l08ld2cy 155 23 , , , cord-346267-l08ld2cy 155 24 even even RB cord-346267-l08ld2cy 155 25 when when WRB cord-346267-l08ld2cy 155 26 the the DT cord-346267-l08ld2cy 155 27 simulated simulated JJ cord-346267-l08ld2cy 155 28 branch branch NN cord-346267-l08ld2cy 155 29 length length NN cord-346267-l08ld2cy 155 30 was be VBD cord-346267-l08ld2cy 155 31 increased increase VBN cord-346267-l08ld2cy 155 32 to to IN cord-346267-l08ld2cy 155 33 100 100 CD cord-346267-l08ld2cy 155 34 ; ; : cord-346267-l08ld2cy 155 35 the the DT cord-346267-l08ld2cy 155 36 branch branch NN cord-346267-l08ld2cy 155 37 length length NN cord-346267-l08ld2cy 155 38 estimate estimate NN cord-346267-l08ld2cy 155 39 was be VBD cord-346267-l08ld2cy 155 40 accurate accurate JJ cord-346267-l08ld2cy 155 41 for for IN cord-346267-l08ld2cy 155 42 up up IN cord-346267-l08ld2cy 155 43 to to TO cord-346267-l08ld2cy 155 44 5 5 CD cord-346267-l08ld2cy 155 45 substitutions substitution NNS cord-346267-l08ld2cy 155 46 / / SYM cord-346267-l08ld2cy 155 47 site site NN cord-346267-l08ld2cy 155 48 . . . cord-346267-l08ld2cy 156 1 For for IN cord-346267-l08ld2cy 156 2 sequences sequence NNS cord-346267-l08ld2cy 156 3 with with IN cord-346267-l08ld2cy 156 4 no no DT cord-346267-l08ld2cy 156 5 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 156 6 substitutions substitution NNS cord-346267-l08ld2cy 156 7 ( ( -LRB- cord-346267-l08ld2cy 156 8 i.e. i.e. FW cord-346267-l08ld2cy 156 9 , , , cord-346267-l08ld2cy 156 10 100 100 CD cord-346267-l08ld2cy 156 11 % % NN cord-346267-l08ld2cy 156 12 conservation conservation NN cord-346267-l08ld2cy 156 13 ) ) -RRB- cord-346267-l08ld2cy 156 14 , , , cord-346267-l08ld2cy 156 15 saturation saturation NN cord-346267-l08ld2cy 156 16 occurred occur VBD cord-346267-l08ld2cy 156 17 much much RB cord-346267-l08ld2cy 156 18 sooner soon RBR cord-346267-l08ld2cy 156 19 , , , cord-346267-l08ld2cy 156 20 around around RB cord-346267-l08ld2cy 156 21 0.25 0.25 CD cord-346267-l08ld2cy 156 22 substitutions substitution NNS cord-346267-l08ld2cy 156 23 / / SYM cord-346267-l08ld2cy 156 24 site site NN cord-346267-l08ld2cy 156 25 , , , cord-346267-l08ld2cy 156 26 and and CC cord-346267-l08ld2cy 156 27 the the DT cord-346267-l08ld2cy 156 28 corresponding corresponding JJ cord-346267-l08ld2cy 156 29 asymptote asymptote NN cord-346267-l08ld2cy 156 30 was be VBD cord-346267-l08ld2cy 156 31 0.36 0.36 CD cord-346267-l08ld2cy 156 32 substitutions substitution NNS cord-346267-l08ld2cy 156 33 / / SYM cord-346267-l08ld2cy 156 34 site site NN cord-346267-l08ld2cy 156 35 . . . cord-346267-l08ld2cy 157 1 The the DT cord-346267-l08ld2cy 157 2 nucleotide nucleotide JJ cord-346267-l08ld2cy 157 3 substitution substitution NN cord-346267-l08ld2cy 157 4 model model NN cord-346267-l08ld2cy 157 5 ( ( -LRB- cord-346267-l08ld2cy 157 6 GTR GTR NNP cord-346267-l08ld2cy 157 7 + + CC cord-346267-l08ld2cy 157 8 Γ γ NN cord-346267-l08ld2cy 157 9 4 4 CD cord-346267-l08ld2cy 157 10 ) ) -RRB- cord-346267-l08ld2cy 157 11 underestimated underestimate VBD cord-346267-l08ld2cy 157 12 the the DT cord-346267-l08ld2cy 157 13 length length NN cord-346267-l08ld2cy 157 14 of of IN cord-346267-l08ld2cy 157 15 branches branch NNS cord-346267-l08ld2cy 157 16 simulated simulate VBN cord-346267-l08ld2cy 157 17 under under IN cord-346267-l08ld2cy 157 18 empirical empirical JJ cord-346267-l08ld2cy 157 19 ( ( -LRB- cord-346267-l08ld2cy 157 20 IFEL IFEL NNP cord-346267-l08ld2cy 157 21 ) ) -RRB- cord-346267-l08ld2cy 157 22 selection selection NN cord-346267-l08ld2cy 157 23 regimes regime NNS cord-346267-l08ld2cy 157 24 inferred infer VBN cord-346267-l08ld2cy 157 25 from from IN cord-346267-l08ld2cy 157 26 the the DT cord-346267-l08ld2cy 157 27 three three CD cord-346267-l08ld2cy 157 28 viral viral JJ cord-346267-l08ld2cy 157 29 data data NN cord-346267-l08ld2cy 157 30 sets set NNS cord-346267-l08ld2cy 157 31 ( ( -LRB- cord-346267-l08ld2cy 157 32 fig fig NNP cord-346267-l08ld2cy 157 33 . . . cord-346267-l08ld2cy 157 34 1B-1D 1b-1d CD cord-346267-l08ld2cy 157 35 ) ) -RRB- cord-346267-l08ld2cy 157 36 ; ; : cord-346267-l08ld2cy 157 37 branch branch NN cord-346267-l08ld2cy 157 38 lengths length NNS cord-346267-l08ld2cy 157 39 for for IN cord-346267-l08ld2cy 157 40 sequences sequence NNS cord-346267-l08ld2cy 157 41 simulated simulate VBN cord-346267-l08ld2cy 157 42 under under IN cord-346267-l08ld2cy 157 43 neutral neutral JJ cord-346267-l08ld2cy 157 44 selection selection NN cord-346267-l08ld2cy 157 45 ( ( -LRB- cord-346267-l08ld2cy 157 46 i.e. i.e. FW cord-346267-l08ld2cy 157 47 , , , cord-346267-l08ld2cy 157 48 α α NN cord-346267-l08ld2cy 157 49 s s NN cord-346267-l08ld2cy 157 50 = = NN cord-346267-l08ld2cy 158 1 β β NNP cord-346267-l08ld2cy 158 2 s s NNP cord-346267-l08ld2cy 158 3 = = SYM cord-346267-l08ld2cy 158 4 1 1 CD cord-346267-l08ld2cy 158 5 ) ) -RRB- cord-346267-l08ld2cy 158 6 were be VBD cord-346267-l08ld2cy 158 7 reliably reliably RB cord-346267-l08ld2cy 158 8 inferred infer VBN cord-346267-l08ld2cy 158 9 by by IN cord-346267-l08ld2cy 158 10 GTR GTR NNP cord-346267-l08ld2cy 158 11 + + CC cord-346267-l08ld2cy 158 12 Γ γ NN cord-346267-l08ld2cy 158 13 4 4 CD cord-346267-l08ld2cy 158 14 . . . cord-346267-l08ld2cy 159 1 The the DT cord-346267-l08ld2cy 159 2 proportion proportion NN cord-346267-l08ld2cy 159 3 of of IN cord-346267-l08ld2cy 159 4 sites site NNS cord-346267-l08ld2cy 159 5 under under IN cord-346267-l08ld2cy 159 6 strong strong JJ cord-346267-l08ld2cy 159 7 negative negative JJ cord-346267-l08ld2cy 159 8 selection selection NN cord-346267-l08ld2cy 159 9 ( ( -LRB- cord-346267-l08ld2cy 159 10 i.e. i.e. FW cord-346267-l08ld2cy 159 11 , , , cord-346267-l08ld2cy 159 12 β β NNP cord-346267-l08ld2cy 159 13 s s NN cord-346267-l08ld2cy 159 14 < < XX cord-346267-l08ld2cy 159 15 α α NNP cord-346267-l08ld2cy 159 16 s s XX cord-346267-l08ld2cy 159 17 with with IN cord-346267-l08ld2cy 159 18 an an DT cord-346267-l08ld2cy 159 19 IFEL IFEL NNP cord-346267-l08ld2cy 159 20 P p NN cord-346267-l08ld2cy 159 21 value value NN cord-346267-l08ld2cy 159 22 0.05 0.05 CD cord-346267-l08ld2cy 159 23 ) ) -RRB- cord-346267-l08ld2cy 160 1 differed differ VBN cord-346267-l08ld2cy 160 2 markedly markedly RB cord-346267-l08ld2cy 160 3 among among IN cord-346267-l08ld2cy 160 4 the the DT cord-346267-l08ld2cy 160 5 three three CD cord-346267-l08ld2cy 160 6 data datum NNS cord-346267-l08ld2cy 160 7 sets set NNS cord-346267-l08ld2cy 160 8 . . . cord-346267-l08ld2cy 161 1 MeV MeV NNP cord-346267-l08ld2cy 161 2 / / SYM cord-346267-l08ld2cy 161 3 RPV RPV NNP cord-346267-l08ld2cy 161 4 / / SYM cord-346267-l08ld2cy 161 5 PPRV PPRV NNP cord-346267-l08ld2cy 161 6 nucleoprotein nucleoprotein NN cord-346267-l08ld2cy 161 7 and and CC cord-346267-l08ld2cy 161 8 EBOV EBOV NNP cord-346267-l08ld2cy 161 9 glycoprotein glycoprotein NN cord-346267-l08ld2cy 161 10 genes gene NNS cord-346267-l08ld2cy 161 11 experienced experience VBD cord-346267-l08ld2cy 161 12 moderate moderate JJ cord-346267-l08ld2cy 161 13 levels level NNS cord-346267-l08ld2cy 161 14 of of IN cord-346267-l08ld2cy 161 15 purifying purify VBG cord-346267-l08ld2cy 161 16 selection selection NN cord-346267-l08ld2cy 161 17 along along IN cord-346267-l08ld2cy 161 18 internal internal JJ cord-346267-l08ld2cy 161 19 branches branch NNS cord-346267-l08ld2cy 161 20 : : : cord-346267-l08ld2cy 161 21 50 50 CD cord-346267-l08ld2cy 161 22 % % NN cord-346267-l08ld2cy 161 23 and and CC cord-346267-l08ld2cy 161 24 45 45 CD cord-346267-l08ld2cy 161 25 % % NN cord-346267-l08ld2cy 161 26 of of IN cord-346267-l08ld2cy 161 27 sites site NNS cord-346267-l08ld2cy 161 28 under under IN cord-346267-l08ld2cy 161 29 strong strong JJ cord-346267-l08ld2cy 161 30 purifying purifying NN cord-346267-l08ld2cy 161 31 selection selection NN cord-346267-l08ld2cy 161 32 , , , cord-346267-l08ld2cy 161 33 respectively respectively RB cord-346267-l08ld2cy 161 34 . . . cord-346267-l08ld2cy 162 1 The the DT cord-346267-l08ld2cy 162 2 AIV AIV NNP cord-346267-l08ld2cy 162 3 neuraminidase neuraminidase NN cord-346267-l08ld2cy 162 4 gene gene NN cord-346267-l08ld2cy 162 5 evolved evolve VBN cord-346267-l08ld2cy 162 6 under under IN cord-346267-l08ld2cy 162 7 much much RB cord-346267-l08ld2cy 162 8 stronger strong JJR cord-346267-l08ld2cy 162 9 selective selective JJ cord-346267-l08ld2cy 162 10 constraints constraint NNS cord-346267-l08ld2cy 162 11 , , , cord-346267-l08ld2cy 162 12 with with IN cord-346267-l08ld2cy 162 13 89 89 CD cord-346267-l08ld2cy 162 14 % % NN cord-346267-l08ld2cy 162 15 of of IN cord-346267-l08ld2cy 162 16 sites site NNS cord-346267-l08ld2cy 162 17 under under IN cord-346267-l08ld2cy 162 18 strong strong JJ cord-346267-l08ld2cy 162 19 purifying purifying NN cord-346267-l08ld2cy 162 20 selection selection NN cord-346267-l08ld2cy 162 21 . . . cord-346267-l08ld2cy 163 1 We -PRON- PRP cord-346267-l08ld2cy 163 2 note note VBP cord-346267-l08ld2cy 163 3 that that IN cord-346267-l08ld2cy 163 4 the the DT cord-346267-l08ld2cy 163 5 power power NN cord-346267-l08ld2cy 163 6 to to TO cord-346267-l08ld2cy 163 7 detect detect VB cord-346267-l08ld2cy 163 8 selection selection NN cord-346267-l08ld2cy 163 9 using use VBG cord-346267-l08ld2cy 163 10 IFEL IFEL NNP cord-346267-l08ld2cy 163 11 increases increase VBZ cord-346267-l08ld2cy 163 12 with with IN cord-346267-l08ld2cy 163 13 the the DT cord-346267-l08ld2cy 163 14 size size NN cord-346267-l08ld2cy 163 15 of of IN cord-346267-l08ld2cy 163 16 the the DT cord-346267-l08ld2cy 163 17 alignment alignment NN cord-346267-l08ld2cy 164 1 ( ( -LRB- cord-346267-l08ld2cy 164 2 Kosakovsky Kosakovsky NNP cord-346267-l08ld2cy 164 3 et et FW cord-346267-l08ld2cy 164 4 al al NNP cord-346267-l08ld2cy 164 5 . . . cord-346267-l08ld2cy 164 6 2006 2006 CD cord-346267-l08ld2cy 164 7 ) ) -RRB- cord-346267-l08ld2cy 165 1 ; ; : cord-346267-l08ld2cy 165 2 hence hence RB cord-346267-l08ld2cy 165 3 , , , cord-346267-l08ld2cy 165 4 we -PRON- PRP cord-346267-l08ld2cy 165 5 would would MD cord-346267-l08ld2cy 165 6 expect expect VB cord-346267-l08ld2cy 165 7 more more JJR cord-346267-l08ld2cy 165 8 sites site NNS cord-346267-l08ld2cy 165 9 evolving evolve VBG cord-346267-l08ld2cy 165 10 under under IN cord-346267-l08ld2cy 165 11 purifying purify VBG cord-346267-l08ld2cy 165 12 selection selection NN cord-346267-l08ld2cy 165 13 to to TO cord-346267-l08ld2cy 165 14 be be VB cord-346267-l08ld2cy 165 15 correctly correctly RB cord-346267-l08ld2cy 165 16 detected detect VBN cord-346267-l08ld2cy 165 17 as as IN cord-346267-l08ld2cy 165 18 such such JJ cord-346267-l08ld2cy 165 19 in in IN cord-346267-l08ld2cy 165 20 the the DT cord-346267-l08ld2cy 165 21 larger large JJR cord-346267-l08ld2cy 165 22 AIV AIV NNP cord-346267-l08ld2cy 165 23 data datum NNS cord-346267-l08ld2cy 165 24 set set VBN cord-346267-l08ld2cy 165 25 . . . cord-346267-l08ld2cy 166 1 The the DT cord-346267-l08ld2cy 166 2 simulated simulated JJ cord-346267-l08ld2cy 166 3 codon codon NN cord-346267-l08ld2cy 166 4 saturation saturation NN cord-346267-l08ld2cy 166 5 curves curve NNS cord-346267-l08ld2cy 166 6 varied varied JJ cord-346267-l08ld2cy 166 7 from from IN cord-346267-l08ld2cy 166 8 virus virus NN cord-346267-l08ld2cy 166 9 to to IN cord-346267-l08ld2cy 166 10 virus virus NN cord-346267-l08ld2cy 166 11 , , , cord-346267-l08ld2cy 166 12 indicating indicate VBG cord-346267-l08ld2cy 166 13 that that IN cord-346267-l08ld2cy 166 14 the the DT cord-346267-l08ld2cy 166 15 particular particular JJ cord-346267-l08ld2cy 166 16 selectiveregime selectiveregime NN cord-346267-l08ld2cy 166 17 , , , cord-346267-l08ld2cy 166 18 amino amino NN cord-346267-l08ld2cy 166 19 acid acid NN cord-346267-l08ld2cy 166 20 composition composition NN cord-346267-l08ld2cy 166 21 , , , cord-346267-l08ld2cy 166 22 and and CC cord-346267-l08ld2cy 166 23 nucleotide nucleotide JJ cord-346267-l08ld2cy 166 24 substitution substitution NN cord-346267-l08ld2cy 166 25 model model NN cord-346267-l08ld2cy 166 26 of of IN cord-346267-l08ld2cy 166 27 each each DT cord-346267-l08ld2cy 166 28 viral viral JJ cord-346267-l08ld2cy 166 29 gene gene NN cord-346267-l08ld2cy 166 30 have have VBP cord-346267-l08ld2cy 166 31 a a DT cord-346267-l08ld2cy 166 32 substantial substantial JJ cord-346267-l08ld2cy 166 33 influence influence NN cord-346267-l08ld2cy 166 34 on on IN cord-346267-l08ld2cy 166 35 the the DT cord-346267-l08ld2cy 166 36 ability ability NN cord-346267-l08ld2cy 166 37 to to TO cord-346267-l08ld2cy 166 38 accurately accurately RB cord-346267-l08ld2cy 166 39 infer infer VB cord-346267-l08ld2cy 166 40 branch branch NN cord-346267-l08ld2cy 166 41 lengths length NNS cord-346267-l08ld2cy 166 42 . . . cord-346267-l08ld2cy 167 1 Although although IN cord-346267-l08ld2cy 167 2 underestimation underestimation NN cord-346267-l08ld2cy 167 3 is be VBZ cord-346267-l08ld2cy 167 4 more more RBR cord-346267-l08ld2cy 167 5 pronounced pronounced JJ cord-346267-l08ld2cy 167 6 along along IN cord-346267-l08ld2cy 167 7 longer long JJR cord-346267-l08ld2cy 167 8 branches branch NNS cord-346267-l08ld2cy 167 9 ( ( -LRB- cord-346267-l08ld2cy 167 10 e.g. e.g. RB cord-346267-l08ld2cy 167 11 , , , cord-346267-l08ld2cy 167 12 about about RB cord-346267-l08ld2cy 167 13 half half PDT cord-346267-l08ld2cy 167 14 the the DT cord-346267-l08ld2cy 167 15 true true JJ cord-346267-l08ld2cy 167 16 length length NN cord-346267-l08ld2cy 167 17 for for IN cord-346267-l08ld2cy 167 18 1 1 CD cord-346267-l08ld2cy 167 19 substitution substitution NN cord-346267-l08ld2cy 167 20 / / SYM cord-346267-l08ld2cy 167 21 site site NN cord-346267-l08ld2cy 167 22 ) ) -RRB- cord-346267-l08ld2cy 167 23 , , , cord-346267-l08ld2cy 167 24 not not RB cord-346267-l08ld2cy 167 25 accounting account VBG cord-346267-l08ld2cy 167 26 for for IN cord-346267-l08ld2cy 167 27 selection selection NN cord-346267-l08ld2cy 167 28 can can MD cord-346267-l08ld2cy 167 29 lead lead VB cord-346267-l08ld2cy 167 30 to to IN cord-346267-l08ld2cy 167 31 underestimation underestimation NN cord-346267-l08ld2cy 167 32 of of IN cord-346267-l08ld2cy 167 33 branch branch NN cord-346267-l08ld2cy 167 34 lengths length NNS cord-346267-l08ld2cy 167 35 even even RB cord-346267-l08ld2cy 167 36 for for IN cord-346267-l08ld2cy 167 37 relatively relatively RB cord-346267-l08ld2cy 167 38 short short JJ cord-346267-l08ld2cy 167 39 branches branch NNS cord-346267-l08ld2cy 167 40 ( ( -LRB- cord-346267-l08ld2cy 167 41 e.g. e.g. RB cord-346267-l08ld2cy 167 42 , , , cord-346267-l08ld2cy 167 43 0.2 0.2 CD cord-346267-l08ld2cy 167 44 - - HYPH cord-346267-l08ld2cy 167 45 0.5 0.5 CD cord-346267-l08ld2cy 167 46 substitutions substitution NNS cord-346267-l08ld2cy 167 47 / / SYM cord-346267-l08ld2cy 167 48 site site NN cord-346267-l08ld2cy 167 49 ) ) -RRB- cord-346267-l08ld2cy 167 50 . . . cord-346267-l08ld2cy 168 1 We -PRON- PRP cord-346267-l08ld2cy 168 2 then then RB cord-346267-l08ld2cy 168 3 inferred infer VBD cord-346267-l08ld2cy 168 4 the the DT cord-346267-l08ld2cy 168 5 length length NN cord-346267-l08ld2cy 168 6 of of IN cord-346267-l08ld2cy 168 7 branches branch NNS cord-346267-l08ld2cy 168 8 simulated simulate VBN cord-346267-l08ld2cy 168 9 under under IN cord-346267-l08ld2cy 168 10 the the DT cord-346267-l08ld2cy 168 11 empirical empirical JJ cord-346267-l08ld2cy 168 12 selective selective JJ cord-346267-l08ld2cy 168 13 regimes regime NNS cord-346267-l08ld2cy 168 14 using use VBG cord-346267-l08ld2cy 168 15 the the DT cord-346267-l08ld2cy 168 16 Dual dual JJ cord-346267-l08ld2cy 168 17 codon codon NN cord-346267-l08ld2cy 168 18 model model NN cord-346267-l08ld2cy 168 19 , , , cord-346267-l08ld2cy 168 20 which which WDT cord-346267-l08ld2cy 168 21 accounts account VBZ cord-346267-l08ld2cy 168 22 for for IN cord-346267-l08ld2cy 168 23 variation variation NN cord-346267-l08ld2cy 168 24 in in IN cord-346267-l08ld2cy 168 25 site site NN cord-346267-l08ld2cy 168 26 - - HYPH cord-346267-l08ld2cy 168 27 to to IN cord-346267-l08ld2cy 168 28 - - HYPH cord-346267-l08ld2cy 168 29 site site NN cord-346267-l08ld2cy 168 30 selective selective JJ cord-346267-l08ld2cy 168 31 pressures pressure NNS cord-346267-l08ld2cy 168 32 ( ( -LRB- cord-346267-l08ld2cy 168 33 fig fig NN cord-346267-l08ld2cy 168 34 . . NNP cord-346267-l08ld2cy 168 35 2 2 CD cord-346267-l08ld2cy 168 36 ) ) -RRB- cord-346267-l08ld2cy 168 37 . . . cord-346267-l08ld2cy 169 1 Although although IN cord-346267-l08ld2cy 169 2 the the DT cord-346267-l08ld2cy 169 3 Dual dual JJ cord-346267-l08ld2cy 169 4 model model NN cord-346267-l08ld2cy 169 5 slightly slightly RB cord-346267-l08ld2cy 169 6 underestimated underestimate VBN cord-346267-l08ld2cy 169 7 lengths length NNS cord-346267-l08ld2cy 169 8 for for IN cord-346267-l08ld2cy 169 9 the the DT cord-346267-l08ld2cy 169 10 longest long JJS cord-346267-l08ld2cy 169 11 simulated simulated JJ cord-346267-l08ld2cy 169 12 branches branch NNS cord-346267-l08ld2cy 170 1 , , , cord-346267-l08ld2cy 170 2 the the DT cord-346267-l08ld2cy 170 3 bias bias NN cord-346267-l08ld2cy 170 4 was be VBD cord-346267-l08ld2cy 170 5 an an DT cord-346267-l08ld2cy 170 6 order order NN cord-346267-l08ld2cy 170 7 of of IN cord-346267-l08ld2cy 170 8 magnitude magnitude NN cord-346267-l08ld2cy 170 9 less less JJR cord-346267-l08ld2cy 170 10 than than IN cord-346267-l08ld2cy 170 11 under under IN cord-346267-l08ld2cy 170 12 GTR GTR NNP cord-346267-l08ld2cy 170 13 + + CC cord-346267-l08ld2cy 170 14 Γ γ NN cord-346267-l08ld2cy 170 15 4 4 CD cord-346267-l08ld2cy 170 16 . . . cord-346267-l08ld2cy 171 1 Furthermore furthermore RB cord-346267-l08ld2cy 171 2 , , , cord-346267-l08ld2cy 171 3 similar similar JJ cord-346267-l08ld2cy 171 4 behavior behavior NN cord-346267-l08ld2cy 171 5 was be VBD cord-346267-l08ld2cy 171 6 observed observe VBN cord-346267-l08ld2cy 171 7 for for IN cord-346267-l08ld2cy 171 8 the the DT cord-346267-l08ld2cy 171 9 longest long JJS cord-346267-l08ld2cy 171 10 branches branch NNS cord-346267-l08ld2cy 171 11 simulated simulate VBN cord-346267-l08ld2cy 171 12 under under IN cord-346267-l08ld2cy 171 13 neutrality neutrality NN cord-346267-l08ld2cy 171 14 ( ( -LRB- cord-346267-l08ld2cy 171 15 α α NN cord-346267-l08ld2cy 171 16 s s NN cord-346267-l08ld2cy 171 17 = = NN cord-346267-l08ld2cy 172 1 β β NNP cord-346267-l08ld2cy 172 2 s s NNP cord-346267-l08ld2cy 172 3 ) ) -RRB- cord-346267-l08ld2cy 172 4 and and CC cord-346267-l08ld2cy 172 5 inferred infer VBD cord-346267-l08ld2cy 172 6 using use VBG cord-346267-l08ld2cy 172 7 GTR GTR NNP cord-346267-l08ld2cy 172 8 + + CC cord-346267-l08ld2cy 172 9 Γ γ NN cord-346267-l08ld2cy 172 10 4 4 CD cord-346267-l08ld2cy 173 1 ( ( -LRB- cord-346267-l08ld2cy 173 2 fig fig NN cord-346267-l08ld2cy 173 3 . . . cord-346267-l08ld2cy 174 1 1B 1B NNP cord-346267-l08ld2cy 174 2 -1D -1D NNP cord-346267-l08ld2cy 174 3 ) ) -RRB- cord-346267-l08ld2cy 174 4 , , . cord-346267-l08ld2cy 175 1 suggesting suggest VBG cord-346267-l08ld2cy 175 2 that that IN cord-346267-l08ld2cy 175 3 both both DT cord-346267-l08ld2cy 175 4 models model NNS cord-346267-l08ld2cy 175 5 perform perform VBP cord-346267-l08ld2cy 175 6 equivalently equivalently RB cord-346267-l08ld2cy 175 7 , , , cord-346267-l08ld2cy 175 8 as as IN cord-346267-l08ld2cy 175 9 expected expect VBN cord-346267-l08ld2cy 175 10 . . . cord-346267-l08ld2cy 176 1 Clearly clearly RB cord-346267-l08ld2cy 176 2 , , , cord-346267-l08ld2cy 176 3 not not RB cord-346267-l08ld2cy 176 4 accounting account VBG cord-346267-l08ld2cy 176 5 for for IN cord-346267-l08ld2cy 176 6 purifying purify VBG cord-346267-l08ld2cy 176 7 selection selection NN cord-346267-l08ld2cy 176 8 can can MD cord-346267-l08ld2cy 176 9 lead lead VB cord-346267-l08ld2cy 176 10 to to IN cord-346267-l08ld2cy 176 11 dramatic dramatic JJ cord-346267-l08ld2cy 176 12 underestimation underestimation NN cord-346267-l08ld2cy 176 13 of of IN cord-346267-l08ld2cy 176 14 branch branch NN cord-346267-l08ld2cy 176 15 lengths length NNS cord-346267-l08ld2cy 176 16 ; ; : cord-346267-l08ld2cy 176 17 this this DT cord-346267-l08ld2cy 176 18 effect effect NN cord-346267-l08ld2cy 176 19 is be VBZ cord-346267-l08ld2cy 176 20 exacerbated exacerbate VBN cord-346267-l08ld2cy 176 21 for for IN cord-346267-l08ld2cy 176 22 longer long JJR cord-346267-l08ld2cy 176 23 branches branch NNS cord-346267-l08ld2cy 176 24 . . . cord-346267-l08ld2cy 177 1 It -PRON- PRP cord-346267-l08ld2cy 177 2 is be VBZ cord-346267-l08ld2cy 177 3 well well RB cord-346267-l08ld2cy 177 4 known know VBN cord-346267-l08ld2cy 177 5 that that IN cord-346267-l08ld2cy 177 6 standard standard JJ cord-346267-l08ld2cy 177 7 nucleotide nucleotide JJ cord-346267-l08ld2cy 177 8 models model NNS cord-346267-l08ld2cy 177 9 differ differ VBP cord-346267-l08ld2cy 177 10 in in IN cord-346267-l08ld2cy 177 11 their -PRON- PRP$ cord-346267-l08ld2cy 177 12 sensitivity sensitivity NN cord-346267-l08ld2cy 177 13 to to IN cord-346267-l08ld2cy 177 14 multiple multiple JJ cord-346267-l08ld2cy 177 15 substitutions substitution NNS cord-346267-l08ld2cy 177 16 at at IN cord-346267-l08ld2cy 177 17 the the DT cord-346267-l08ld2cy 177 18 same same JJ cord-346267-l08ld2cy 177 19 site site NN cord-346267-l08ld2cy 177 20 ( ( -LRB- cord-346267-l08ld2cy 177 21 Sullivan Sullivan NNP cord-346267-l08ld2cy 177 22 and and CC cord-346267-l08ld2cy 177 23 Joyce Joyce NNP cord-346267-l08ld2cy 177 24 2005 2005 CD cord-346267-l08ld2cy 177 25 ) ) -RRB- cord-346267-l08ld2cy 177 26 . . . cord-346267-l08ld2cy 178 1 We -PRON- PRP cord-346267-l08ld2cy 178 2 explored explore VBD cord-346267-l08ld2cy 178 3 how how WRB cord-346267-l08ld2cy 178 4 various various JJ cord-346267-l08ld2cy 178 5 methods method NNS cord-346267-l08ld2cy 178 6 of of IN cord-346267-l08ld2cy 178 7 modeling model VBG cord-346267-l08ld2cy 178 8 rate rate NN cord-346267-l08ld2cy 178 9 variation variation NN cord-346267-l08ld2cy 178 10 affected affect VBN cord-346267-l08ld2cy 178 11 branch branch NN cord-346267-l08ld2cy 178 12 length length NN cord-346267-l08ld2cy 178 13 inference inference NN cord-346267-l08ld2cy 178 14 in in IN cord-346267-l08ld2cy 178 15 our -PRON- PRP$ cord-346267-l08ld2cy 178 16 three three CD cord-346267-l08ld2cy 178 17 viral viral JJ cord-346267-l08ld2cy 178 18 data data NN cord-346267-l08ld2cy 178 19 sets set NNS cord-346267-l08ld2cy 178 20 , , , cord-346267-l08ld2cy 178 21 using use VBG cord-346267-l08ld2cy 178 22 a a DT cord-346267-l08ld2cy 178 23 fixed fix VBN cord-346267-l08ld2cy 178 24 MCC MCC NNP cord-346267-l08ld2cy 178 25 tree tree NN cord-346267-l08ld2cy 178 26 from from IN cord-346267-l08ld2cy 178 27 the the DT cord-346267-l08ld2cy 178 28 GTR GTR NNP cord-346267-l08ld2cy 178 29 + + SYM cord-346267-l08ld2cy 178 30 Γ γ NN cord-346267-l08ld2cy 178 31 4 4 CD cord-346267-l08ld2cy 178 32 BMCMC BMCMC NNP cord-346267-l08ld2cy 178 33 analyses analysis NNS cord-346267-l08ld2cy 178 34 . . . cord-346267-l08ld2cy 179 1 First first RB cord-346267-l08ld2cy 179 2 , , , cord-346267-l08ld2cy 179 3 we -PRON- PRP cord-346267-l08ld2cy 179 4 examined examine VBD cord-346267-l08ld2cy 179 5 two two CD cord-346267-l08ld2cy 179 6 extremes extreme NNS cord-346267-l08ld2cy 179 7 among among IN cord-346267-l08ld2cy 179 8 nucleotide nucleotide JJ cord-346267-l08ld2cy 179 9 substitution substitution NN cord-346267-l08ld2cy 179 10 models model NNS cord-346267-l08ld2cy 179 11 that that WDT cord-346267-l08ld2cy 179 12 do do VBP cord-346267-l08ld2cy 179 13 not not RB cord-346267-l08ld2cy 179 14 adjust adjust VB cord-346267-l08ld2cy 179 15 for for IN cord-346267-l08ld2cy 179 16 rate rate NN cord-346267-l08ld2cy 179 17 variation variation NN cord-346267-l08ld2cy 179 18 across across IN cord-346267-l08ld2cy 179 19 sites site NNS cord-346267-l08ld2cy 179 20 : : : cord-346267-l08ld2cy 179 21 JC69 JC69 NNP cord-346267-l08ld2cy 179 22 ( ( -LRB- cord-346267-l08ld2cy 179 23 Jukes Jukes NNP cord-346267-l08ld2cy 179 24 and and CC cord-346267-l08ld2cy 179 25 Cantor Cantor NNP cord-346267-l08ld2cy 179 26 1969 1969 CD cord-346267-l08ld2cy 179 27 ) ) -RRB- cord-346267-l08ld2cy 179 28 and and CC cord-346267-l08ld2cy 179 29 GTR GTR NNP cord-346267-l08ld2cy 179 30 . . . cord-346267-l08ld2cy 180 1 JC69 JC69 NNP cord-346267-l08ld2cy 180 2 assumes assume VBZ cord-346267-l08ld2cy 180 3 a a DT cord-346267-l08ld2cy 180 4 single single JJ cord-346267-l08ld2cy 180 5 substitution substitution NN cord-346267-l08ld2cy 180 6 rate rate NN cord-346267-l08ld2cy 180 7 and and CC cord-346267-l08ld2cy 180 8 equal equal JJ cord-346267-l08ld2cy 180 9 base base NN cord-346267-l08ld2cy 180 10 frequencies frequency NNS cord-346267-l08ld2cy 180 11 , , , cord-346267-l08ld2cy 180 12 whereas whereas IN cord-346267-l08ld2cy 180 13 GTR GTR NNP cord-346267-l08ld2cy 180 14 allows allow VBZ cord-346267-l08ld2cy 180 15 each each DT cord-346267-l08ld2cy 180 16 class class NN cord-346267-l08ld2cy 180 17 of of IN cord-346267-l08ld2cy 180 18 nucleotide nucleotide JJ cord-346267-l08ld2cy 180 19 substitution substitution NN cord-346267-l08ld2cy 180 20 to to TO cord-346267-l08ld2cy 180 21 occur occur VB cord-346267-l08ld2cy 180 22 at at IN cord-346267-l08ld2cy 180 23 a a DT cord-346267-l08ld2cy 180 24 unique unique JJ cord-346267-l08ld2cy 180 25 rate rate NN cord-346267-l08ld2cy 180 26 and and CC cord-346267-l08ld2cy 180 27 estimates estimate VBZ cord-346267-l08ld2cy 180 28 base base VBP cord-346267-l08ld2cy 180 29 frequencies frequency NNS cord-346267-l08ld2cy 180 30 from from IN cord-346267-l08ld2cy 180 31 the the DT cord-346267-l08ld2cy 180 32 data datum NNS cord-346267-l08ld2cy 180 33 . . . cord-346267-l08ld2cy 181 1 As as IN cord-346267-l08ld2cy 181 2 expected expect VBN cord-346267-l08ld2cy 181 3 , , , cord-346267-l08ld2cy 181 4 failure failure NN cord-346267-l08ld2cy 181 5 to to TO cord-346267-l08ld2cy 181 6 account account VB cord-346267-l08ld2cy 181 7 for for IN cord-346267-l08ld2cy 181 8 site site NN cord-346267-l08ld2cy 181 9 - - HYPH cord-346267-l08ld2cy 181 10 to to IN cord-346267-l08ld2cy 181 11 - - HYPH cord-346267-l08ld2cy 181 12 site site NN cord-346267-l08ld2cy 181 13 rate rate NN cord-346267-l08ld2cy 181 14 variation variation NN cord-346267-l08ld2cy 181 15 led lead VBD cord-346267-l08ld2cy 181 16 to to IN cord-346267-l08ld2cy 181 17 severe severe JJ cord-346267-l08ld2cy 181 18 underestimation underestimation NN cord-346267-l08ld2cy 181 19 of of IN cord-346267-l08ld2cy 181 20 longer long JJR cord-346267-l08ld2cy 181 21 branches branch NNS cord-346267-l08ld2cy 181 22 in in IN cord-346267-l08ld2cy 181 23 all all DT cord-346267-l08ld2cy 181 24 three three CD cord-346267-l08ld2cy 181 25 viral viral JJ cord-346267-l08ld2cy 181 26 data data NN cord-346267-l08ld2cy 181 27 sets set NNS cord-346267-l08ld2cy 181 28 ( ( -LRB- cord-346267-l08ld2cy 181 29 JC69 JC69 NNP cord-346267-l08ld2cy 181 30 or or CC cord-346267-l08ld2cy 181 31 GTR GTR NNP cord-346267-l08ld2cy 181 32 vs. vs. IN cord-346267-l08ld2cy 181 33 GTR GTR NNP cord-346267-l08ld2cy 181 34 + + CC cord-346267-l08ld2cy 181 35 Γ γ NN cord-346267-l08ld2cy 181 36 4 4 CD cord-346267-l08ld2cy 181 37 , , , cord-346267-l08ld2cy 181 38 fig fig NN cord-346267-l08ld2cy 181 39 . . . cord-346267-l08ld2cy 182 1 3 3 LS cord-346267-l08ld2cy 183 1 the the DT cord-346267-l08ld2cy 183 2 importance importance NN cord-346267-l08ld2cy 183 3 of of IN cord-346267-l08ld2cy 183 4 modeling model VBG cord-346267-l08ld2cy 183 5 site site NN cord-346267-l08ld2cy 183 6 - - HYPH cord-346267-l08ld2cy 183 7 to to IN cord-346267-l08ld2cy 183 8 - - HYPH cord-346267-l08ld2cy 183 9 site site NN cord-346267-l08ld2cy 183 10 rate rate NN cord-346267-l08ld2cy 183 11 variation variation NN cord-346267-l08ld2cy 183 12 to to IN cord-346267-l08ld2cy 183 13 accurate accurate JJ cord-346267-l08ld2cy 183 14 inference inference NN cord-346267-l08ld2cy 183 15 . . . cord-346267-l08ld2cy 184 1 Allowing allow VBG cord-346267-l08ld2cy 184 2 for for IN cord-346267-l08ld2cy 184 3 multiple multiple JJ cord-346267-l08ld2cy 184 4 substitution substitution NN cord-346267-l08ld2cy 184 5 rates rate NNS cord-346267-l08ld2cy 184 6 and and CC cord-346267-l08ld2cy 184 7 empirical empirical JJ cord-346267-l08ld2cy 184 8 base base NN cord-346267-l08ld2cy 184 9 frequencies frequency NNS cord-346267-l08ld2cy 184 10 had have VBD cord-346267-l08ld2cy 184 11 a a DT cord-346267-l08ld2cy 184 12 negligible negligible JJ cord-346267-l08ld2cy 184 13 impact impact NN cord-346267-l08ld2cy 184 14 on on IN cord-346267-l08ld2cy 184 15 branch branch NN cord-346267-l08ld2cy 184 16 length length NN cord-346267-l08ld2cy 184 17 estimation estimation NN cord-346267-l08ld2cy 184 18 ( ( -LRB- cord-346267-l08ld2cy 184 19 fig fig NN cord-346267-l08ld2cy 184 20 . . . cord-346267-l08ld2cy 184 21 3 3 CD cord-346267-l08ld2cy 184 22 ) ) -RRB- cord-346267-l08ld2cy 184 23 . . . cord-346267-l08ld2cy 185 1 Underestimation underestimation NN cord-346267-l08ld2cy 185 2 of of IN cord-346267-l08ld2cy 185 3 long long JJ cord-346267-l08ld2cy 185 4 branches branch NNS cord-346267-l08ld2cy 185 5 likely likely RB cord-346267-l08ld2cy 185 6 explains explain VBZ cord-346267-l08ld2cy 185 7 why why WRB cord-346267-l08ld2cy 185 8 the the DT cord-346267-l08ld2cy 185 9 GTR GTR NNP cord-346267-l08ld2cy 185 10 model model NN cord-346267-l08ld2cy 185 11 inferred infer VBD cord-346267-l08ld2cy 185 12 younger young JJR cord-346267-l08ld2cy 185 13 root root VB cord-346267-l08ld2cy 185 14 tMRCAs tmrcas CD cord-346267-l08ld2cy 185 15 for for IN cord-346267-l08ld2cy 185 16 all all PDT cord-346267-l08ld2cy 185 17 all all DT cord-346267-l08ld2cy 185 18 three three CD cord-346267-l08ld2cy 185 19 viral viral JJ cord-346267-l08ld2cy 185 20 data data NN cord-346267-l08ld2cy 185 21 sets set NNS cord-346267-l08ld2cy 185 22 ( ( -LRB- cord-346267-l08ld2cy 185 23 table table NN cord-346267-l08ld2cy 185 24 1 1 CD cord-346267-l08ld2cy 185 25 ) ) -RRB- cord-346267-l08ld2cy 185 26 . . . cord-346267-l08ld2cy 186 1 We -PRON- PRP cord-346267-l08ld2cy 186 2 hypothesized hypothesize VBD cord-346267-l08ld2cy 186 3 that that IN cord-346267-l08ld2cy 186 4 codon codon NN cord-346267-l08ld2cy 186 5 substitution substitution NN cord-346267-l08ld2cy 186 6 models model NNS cord-346267-l08ld2cy 186 7 , , , cord-346267-l08ld2cy 186 8 which which WDT cord-346267-l08ld2cy 186 9 explicitly explicitly RB cord-346267-l08ld2cy 186 10 account account VBP cord-346267-l08ld2cy 186 11 for for IN cord-346267-l08ld2cy 186 12 the the DT cord-346267-l08ld2cy 186 13 differences difference NNS cord-346267-l08ld2cy 186 14 between between IN cord-346267-l08ld2cy 186 15 synonymous synonymous JJ cord-346267-l08ld2cy 186 16 and and CC cord-346267-l08ld2cy 186 17 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 186 18 substitutions substitution NNS cord-346267-l08ld2cy 186 19 , , , cord-346267-l08ld2cy 186 20 would would MD cord-346267-l08ld2cy 186 21 permit permit VB cord-346267-l08ld2cy 186 22 a a DT cord-346267-l08ld2cy 186 23 more more RBR cord-346267-l08ld2cy 186 24 accurate accurate JJ cord-346267-l08ld2cy 186 25 estimation estimation NN cord-346267-l08ld2cy 186 26 of of IN cord-346267-l08ld2cy 186 27 sequence sequence NN cord-346267-l08ld2cy 186 28 divergence divergence NN cord-346267-l08ld2cy 186 29 well well RB cord-346267-l08ld2cy 186 30 past past IN cord-346267-l08ld2cy 186 31 the the DT cord-346267-l08ld2cy 186 32 point point NN cord-346267-l08ld2cy 186 33 a a DT cord-346267-l08ld2cy 186 34 standard standard JJ cord-346267-l08ld2cy 186 35 nucleotide nucleotide JJ cord-346267-l08ld2cy 186 36 model model NN cord-346267-l08ld2cy 186 37 would would MD cord-346267-l08ld2cy 186 38 reach reach VB cord-346267-l08ld2cy 186 39 saturation saturation NN cord-346267-l08ld2cy 186 40 . . . cord-346267-l08ld2cy 187 1 Therefore therefore RB cord-346267-l08ld2cy 187 2 , , , cord-346267-l08ld2cy 187 3 it -PRON- PRP cord-346267-l08ld2cy 187 4 may may MD cord-346267-l08ld2cy 187 5 be be VB cord-346267-l08ld2cy 187 6 possible possible JJ cord-346267-l08ld2cy 187 7 to to TO cord-346267-l08ld2cy 187 8 use use VB cord-346267-l08ld2cy 187 9 RNA RNA NNP cord-346267-l08ld2cy 187 10 virus virus NN cord-346267-l08ld2cy 187 11 genes gene NNS cord-346267-l08ld2cy 187 12 , , , cord-346267-l08ld2cy 187 13 which which WDT cord-346267-l08ld2cy 187 14 evolve evolve VBP cord-346267-l08ld2cy 187 15 extremely extremely RB cord-346267-l08ld2cy 187 16 rapidly rapidly RB cord-346267-l08ld2cy 187 17 ( ( -LRB- cord-346267-l08ld2cy 187 18 e.g. e.g. RB cord-346267-l08ld2cy 187 19 , , , cord-346267-l08ld2cy 187 20 0.0001 0.0001 CD cord-346267-l08ld2cy 187 21 - - HYPH cord-346267-l08ld2cy 187 22 0.001 0.001 CD cord-346267-l08ld2cy 187 23 substitutions substitution NNS cord-346267-l08ld2cy 187 24 / / SYM cord-346267-l08ld2cy 187 25 site site NN cord-346267-l08ld2cy 187 26 / / SYM cord-346267-l08ld2cy 187 27 year year NN cord-346267-l08ld2cy 187 28 ) ) -RRB- cord-346267-l08ld2cy 187 29 , , , cord-346267-l08ld2cy 187 30 to to TO cord-346267-l08ld2cy 187 31 estimate estimate VB cord-346267-l08ld2cy 187 32 ancient ancient JJ cord-346267-l08ld2cy 187 33 viral viral JJ cord-346267-l08ld2cy 187 34 divergence divergence NN cord-346267-l08ld2cy 187 35 events event NNS cord-346267-l08ld2cy 187 36 using use VBG cord-346267-l08ld2cy 187 37 codon codon NN cord-346267-l08ld2cy 187 38 substitution substitution NN cord-346267-l08ld2cy 187 39 models model NNS cord-346267-l08ld2cy 187 40 . . . cord-346267-l08ld2cy 188 1 A a DT cord-346267-l08ld2cy 188 2 crude crude JJ cord-346267-l08ld2cy 188 3 approximation approximation NN cord-346267-l08ld2cy 188 4 of of IN cord-346267-l08ld2cy 188 5 a a DT cord-346267-l08ld2cy 188 6 codon codon NN cord-346267-l08ld2cy 188 7 model model NN cord-346267-l08ld2cy 188 8 , , , cord-346267-l08ld2cy 188 9 SRD06 SRD06 NNP cord-346267-l08ld2cy 188 10 , , , cord-346267-l08ld2cy 188 11 produced produce VBN cord-346267-l08ld2cy 188 12 branch branch NN cord-346267-l08ld2cy 188 13 lengths length NNS cord-346267-l08ld2cy 188 14 that that WDT cord-346267-l08ld2cy 188 15 were be VBD cord-346267-l08ld2cy 188 16 essentially essentially RB cord-346267-l08ld2cy 188 17 the the DT cord-346267-l08ld2cy 188 18 same same JJ cord-346267-l08ld2cy 188 19 as as IN cord-346267-l08ld2cy 188 20 those those DT cord-346267-l08ld2cy 188 21 from from IN cord-346267-l08ld2cy 188 22 the the DT cord-346267-l08ld2cy 188 23 GTR GTR NNP cord-346267-l08ld2cy 188 24 + + SYM cord-346267-l08ld2cy 188 25 Γ γ NN cord-346267-l08ld2cy 188 26 4 4 CD cord-346267-l08ld2cy 188 27 model model NN cord-346267-l08ld2cy 188 28 ( ( -LRB- cord-346267-l08ld2cy 188 29 fig fig NN cord-346267-l08ld2cy 188 30 . . . cord-346267-l08ld2cy 188 31 3 3 LS cord-346267-l08ld2cy 188 32 ) ) -RRB- cord-346267-l08ld2cy 188 33 , , , cord-346267-l08ld2cy 188 34 which which WDT cord-346267-l08ld2cy 188 35 may may MD cord-346267-l08ld2cy 188 36 explain explain VB cord-346267-l08ld2cy 188 37 why why WRB cord-346267-l08ld2cy 188 38 BMCMC BMCMC NNP cord-346267-l08ld2cy 188 39 tMRCA tmrca XX cord-346267-l08ld2cy 188 40 inference inference NN cord-346267-l08ld2cy 188 41 using use VBG cord-346267-l08ld2cy 188 42 these these DT cord-346267-l08ld2cy 188 43 two two CD cord-346267-l08ld2cy 188 44 models model NNS cord-346267-l08ld2cy 188 45 was be VBD cord-346267-l08ld2cy 188 46 so so RB cord-346267-l08ld2cy 188 47 similar similar JJ cord-346267-l08ld2cy 188 48 . . . cord-346267-l08ld2cy 189 1 The the DT cord-346267-l08ld2cy 189 2 inference inference NN cord-346267-l08ld2cy 189 3 under under IN cord-346267-l08ld2cy 189 4 the the DT cord-346267-l08ld2cy 189 5 GY94 GY94 NNP cord-346267-l08ld2cy 189 6 + + CC cord-346267-l08ld2cy 189 7 Γ γ NN cord-346267-l08ld2cy 189 8 4 4 CD cord-346267-l08ld2cy 189 9 codon codon NN cord-346267-l08ld2cy 189 10 model model NN cord-346267-l08ld2cy 189 11 resulted result VBD cord-346267-l08ld2cy 189 12 in in IN cord-346267-l08ld2cy 189 13 longer long JJR cord-346267-l08ld2cy 189 14 internal internal JJ cord-346267-l08ld2cy 189 15 branch branch NN cord-346267-l08ld2cy 189 16 lengths length NNS cord-346267-l08ld2cy 189 17 only only RB cord-346267-l08ld2cy 189 18 for for IN cord-346267-l08ld2cy 189 19 EBOV EBOV NNP cord-346267-l08ld2cy 189 20 ( ( -LRB- cord-346267-l08ld2cy 190 1 fig fig NNP cord-346267-l08ld2cy 190 2 . . . cord-346267-l08ld2cy 190 3 3 3 LS cord-346267-l08ld2cy 190 4 ) ) -RRB- cord-346267-l08ld2cy 190 5 , , , cord-346267-l08ld2cy 190 6 which which WDT cord-346267-l08ld2cy 190 7 was be VBD cord-346267-l08ld2cy 190 8 in in IN cord-346267-l08ld2cy 190 9 agreement agreement NN cord-346267-l08ld2cy 190 10 with with IN cord-346267-l08ld2cy 190 11 our -PRON- PRP$ cord-346267-l08ld2cy 190 12 BMCMC BMCMC NNP cord-346267-l08ld2cy 190 13 tMRCA tmrca NN cord-346267-l08ld2cy 190 14 inference inference NN cord-346267-l08ld2cy 190 15 using use VBG cord-346267-l08ld2cy 190 16 this this DT cord-346267-l08ld2cy 190 17 model model NN cord-346267-l08ld2cy 190 18 . . . cord-346267-l08ld2cy 191 1 Based base VBN cord-346267-l08ld2cy 191 2 on on IN cord-346267-l08ld2cy 191 3 the the DT cord-346267-l08ld2cy 191 4 simulations simulation NNS cord-346267-l08ld2cy 191 5 along along IN cord-346267-l08ld2cy 191 6 a a DT cord-346267-l08ld2cy 191 7 single single JJ cord-346267-l08ld2cy 191 8 branch branch NN cord-346267-l08ld2cy 191 9 , , , cord-346267-l08ld2cy 191 10 we -PRON- PRP cord-346267-l08ld2cy 191 11 anticipated anticipate VBD cord-346267-l08ld2cy 191 12 that that IN cord-346267-l08ld2cy 191 13 longer long JJR cord-346267-l08ld2cy 191 14 branches branch NNS cord-346267-l08ld2cy 191 15 would would MD cord-346267-l08ld2cy 191 16 be be VB cord-346267-l08ld2cy 191 17 disproportionately disproportionately RB cord-346267-l08ld2cy 191 18 affected affect VBN cord-346267-l08ld2cy 191 19 by by IN cord-346267-l08ld2cy 191 20 site site NN cord-346267-l08ld2cy 191 21 - - HYPH cord-346267-l08ld2cy 191 22 to to IN cord-346267-l08ld2cy 191 23 - - HYPH cord-346267-l08ld2cy 191 24 site site NN cord-346267-l08ld2cy 191 25 variation variation NN cord-346267-l08ld2cy 191 26 in in IN cord-346267-l08ld2cy 191 27 selection selection NN cord-346267-l08ld2cy 191 28 pressures pressure NNS cord-346267-l08ld2cy 191 29 . . . cord-346267-l08ld2cy 192 1 Therefore therefore RB cord-346267-l08ld2cy 192 2 , , , cord-346267-l08ld2cy 192 3 we -PRON- PRP cord-346267-l08ld2cy 192 4 investigated investigate VBD cord-346267-l08ld2cy 192 5 how how WRB cord-346267-l08ld2cy 192 6 extensions extension NNS cord-346267-l08ld2cy 192 7 of of IN cord-346267-l08ld2cy 192 8 the the DT cord-346267-l08ld2cy 192 9 MG94 MG94 NNP cord-346267-l08ld2cy 192 10 model model NN cord-346267-l08ld2cy 192 11 that that WDT cord-346267-l08ld2cy 192 12 incorporate incorporate VBP cord-346267-l08ld2cy 192 13 β β XX cord-346267-l08ld2cy 192 14 , , , cord-346267-l08ld2cy 192 15 α α NN cord-346267-l08ld2cy 192 16 , , , cord-346267-l08ld2cy 192 17 and and CC cord-346267-l08ld2cy 192 18 lineage lineage NN cord-346267-l08ld2cy 192 19 - - HYPH cord-346267-l08ld2cy 192 20 specific specific JJ cord-346267-l08ld2cy 192 21 rate rate NN cord-346267-l08ld2cy 192 22 variation variation NN cord-346267-l08ld2cy 192 23 affected affect VBN cord-346267-l08ld2cy 192 24 branch branch NN cord-346267-l08ld2cy 192 25 length length NN cord-346267-l08ld2cy 192 26 inference inference NN cord-346267-l08ld2cy 192 27 ( ( -LRB- cord-346267-l08ld2cy 192 28 table table NN cord-346267-l08ld2cy 192 29 2 2 CD cord-346267-l08ld2cy 192 30 ) ) -RRB- cord-346267-l08ld2cy 192 31 . . . cord-346267-l08ld2cy 193 1 For for IN cord-346267-l08ld2cy 193 2 MeV MeV NNP cord-346267-l08ld2cy 193 3 and and CC cord-346267-l08ld2cy 193 4 AIV AIV NNP cord-346267-l08ld2cy 193 5 , , , cord-346267-l08ld2cy 193 6 the the DT cord-346267-l08ld2cy 193 7 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 193 8 ( ( -LRB- cord-346267-l08ld2cy 193 9 two two CD cord-346267-l08ld2cy 193 10 - - HYPH cord-346267-l08ld2cy 193 11 rate rate NN cord-346267-l08ld2cy 193 12 ) ) -RRB- cord-346267-l08ld2cy 193 13 model model NN cord-346267-l08ld2cy 193 14 that that WDT cord-346267-l08ld2cy 193 15 allowed allow VBD cord-346267-l08ld2cy 193 16 longer long JJR cord-346267-l08ld2cy 193 17 internal internal JJ cord-346267-l08ld2cy 193 18 branches branch NNS cord-346267-l08ld2cy 193 19 to to TO cord-346267-l08ld2cy 193 20 share share VB cord-346267-l08ld2cy 193 21 their -PRON- PRP$ cord-346267-l08ld2cy 193 22 own own JJ cord-346267-l08ld2cy 193 23 ratio ratio NN cord-346267-l08ld2cy 194 1 E e LS cord-346267-l08ld2cy 194 2 [ [ -LRB- cord-346267-l08ld2cy 194 3 β]/E β]/e XX cord-346267-l08ld2cy 194 4 [ [ -LRB- cord-346267-l08ld2cy 194 5 α α NN cord-346267-l08ld2cy 194 6 ] ] -RRB- cord-346267-l08ld2cy 194 7 provided provide VBD cord-346267-l08ld2cy 194 8 the the DT cord-346267-l08ld2cy 194 9 best good JJS cord-346267-l08ld2cy 194 10 fit fit NN cord-346267-l08ld2cy 194 11 to to IN cord-346267-l08ld2cy 194 12 the the DT cord-346267-l08ld2cy 194 13 data datum NNS cord-346267-l08ld2cy 194 14 . . . cord-346267-l08ld2cy 195 1 For for IN cord-346267-l08ld2cy 195 2 EBOV EBOV NNP cord-346267-l08ld2cy 195 3 , , , cord-346267-l08ld2cy 195 4 a a DT cord-346267-l08ld2cy 195 5 more more RBR cord-346267-l08ld2cy 195 6 complicated complicated JJ cord-346267-l08ld2cy 195 7 ( ( -LRB- cord-346267-l08ld2cy 195 8 eight eight CD cord-346267-l08ld2cy 195 9 - - HYPH cord-346267-l08ld2cy 195 10 rate rate NN cord-346267-l08ld2cy 195 11 ) ) -RRB- cord-346267-l08ld2cy 195 12 model model NN cord-346267-l08ld2cy 195 13 , , , cord-346267-l08ld2cy 195 14 in in IN cord-346267-l08ld2cy 195 15 which which WDT cord-346267-l08ld2cy 195 16 each each DT cord-346267-l08ld2cy 195 17 intersubtype intersubtype NNP cord-346267-l08ld2cy 195 18 branch branch NN cord-346267-l08ld2cy 195 19 and and CC cord-346267-l08ld2cy 195 20 the the DT cord-346267-l08ld2cy 195 21 long long JJ cord-346267-l08ld2cy 195 22 terminal terminal NN cord-346267-l08ld2cy 195 23 branches branch NNS cord-346267-l08ld2cy 195 24 leading lead VBG cord-346267-l08ld2cy 195 25 to to IN cord-346267-l08ld2cy 195 26 Côte Côte NNP cord-346267-l08ld2cy 195 27 d'Ivoire d'Ivoire NNP cord-346267-l08ld2cy 195 28 and and CC cord-346267-l08ld2cy 195 29 Bundibugyo Bundibugyo NNP cord-346267-l08ld2cy 195 30 had have VBD cord-346267-l08ld2cy 195 31 their -PRON- PRP$ cord-346267-l08ld2cy 195 32 own own JJ cord-346267-l08ld2cy 195 33 rates rate NNS cord-346267-l08ld2cy 195 34 , , , cord-346267-l08ld2cy 195 35 was be VBD cord-346267-l08ld2cy 195 36 fit fit JJ cord-346267-l08ld2cy 195 37 because because IN cord-346267-l08ld2cy 195 38 E e NN cord-346267-l08ld2cy 195 39 [ [ -LRB- cord-346267-l08ld2cy 195 40 β]/E β]/e XX cord-346267-l08ld2cy 195 41 [ [ -LRB- cord-346267-l08ld2cy 195 42 α α NN cord-346267-l08ld2cy 195 43 ] ] -RRB- cord-346267-l08ld2cy 195 44 varied vary VBN cord-346267-l08ld2cy 195 45 dramatically dramatically RB cord-346267-l08ld2cy 195 46 among among IN cord-346267-l08ld2cy 195 47 the the DT cord-346267-l08ld2cy 195 48 intersubtype intersubtype NNP cord-346267-l08ld2cy 195 49 branches branch NNS cord-346267-l08ld2cy 195 50 . . . cord-346267-l08ld2cy 196 1 The the DT cord-346267-l08ld2cy 196 2 inclusion inclusion NN cord-346267-l08ld2cy 196 3 of of IN cord-346267-l08ld2cy 196 4 variation variation NN cord-346267-l08ld2cy 196 5 in in IN cord-346267-l08ld2cy 196 6 β β NN cord-346267-l08ld2cy 196 7 and and CC cord-346267-l08ld2cy 196 8 α α NN cord-346267-l08ld2cy 196 9 in in IN cord-346267-l08ld2cy 196 10 the the DT cord-346267-l08ld2cy 196 11 Nonsynonymous Nonsynonymous NNP cord-346267-l08ld2cy 196 12 and and CC cord-346267-l08ld2cy 196 13 Dual dual JJ cord-346267-l08ld2cy 196 14 models model NNS cord-346267-l08ld2cy 196 15 resulted result VBD cord-346267-l08ld2cy 196 16 in in IN cord-346267-l08ld2cy 196 17 notable notable JJ cord-346267-l08ld2cy 196 18 increases increase NNS cord-346267-l08ld2cy 196 19 in in IN cord-346267-l08ld2cy 196 20 the the DT cord-346267-l08ld2cy 196 21 cumulative cumulative JJ cord-346267-l08ld2cy 196 22 length length NN cord-346267-l08ld2cy 196 23 of of IN cord-346267-l08ld2cy 196 24 deep deep JJ cord-346267-l08ld2cy 196 25 internal internal JJ cord-346267-l08ld2cy 196 26 branches branch NNS cord-346267-l08ld2cy 196 27 ( ( -LRB- cord-346267-l08ld2cy 196 28 table table NN cord-346267-l08ld2cy 196 29 2 2 CD cord-346267-l08ld2cy 196 30 ; ; , cord-346267-l08ld2cy 196 31 supplementary supplementary JJ cord-346267-l08ld2cy 196 32 fig fig NN cord-346267-l08ld2cy 196 33 . . . cord-346267-l08ld2cy 197 1 S1 S1 NNP cord-346267-l08ld2cy 197 2 , , , cord-346267-l08ld2cy 197 3 Supplementary Supplementary NNP cord-346267-l08ld2cy 197 4 Material Material NNP cord-346267-l08ld2cy 197 5 online online RB cord-346267-l08ld2cy 197 6 ) ) -RRB- cord-346267-l08ld2cy 197 7 . . . cord-346267-l08ld2cy 198 1 In in IN cord-346267-l08ld2cy 198 2 all all DT cord-346267-l08ld2cy 198 3 three three CD cord-346267-l08ld2cy 198 4 viral viral JJ cord-346267-l08ld2cy 198 5 data data NN cord-346267-l08ld2cy 198 6 sets set NNS cord-346267-l08ld2cy 198 7 , , , cord-346267-l08ld2cy 198 8 however however RB cord-346267-l08ld2cy 198 9 , , , cord-346267-l08ld2cy 198 10 the the DT cord-346267-l08ld2cy 198 11 most most RBS cord-346267-l08ld2cy 198 12 substantial substantial JJ cord-346267-l08ld2cy 198 13 differences difference NNS cord-346267-l08ld2cy 198 14 were be VBD cord-346267-l08ld2cy 198 15 seen see VBN cord-346267-l08ld2cy 198 16 with with IN cord-346267-l08ld2cy 198 17 the the DT cord-346267-l08ld2cy 198 18 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 198 19 model model NN cord-346267-l08ld2cy 198 20 , , , cord-346267-l08ld2cy 198 21 which which WDT cord-346267-l08ld2cy 198 22 produced produce VBD cord-346267-l08ld2cy 198 23 lengths length NNS cord-346267-l08ld2cy 198 24 for for IN cord-346267-l08ld2cy 198 25 long long JJ cord-346267-l08ld2cy 198 26 internal internal JJ cord-346267-l08ld2cy 198 27 branches branch NNS cord-346267-l08ld2cy 198 28 that that WDT cord-346267-l08ld2cy 198 29 were be VBD cord-346267-l08ld2cy 198 30 substantially substantially RB cord-346267-l08ld2cy 198 31 greater great JJR cord-346267-l08ld2cy 198 32 than than IN cord-346267-l08ld2cy 198 33 those those DT cord-346267-l08ld2cy 198 34 inferred infer VBN cord-346267-l08ld2cy 198 35 under under IN cord-346267-l08ld2cy 198 36 GTR GTR NNP cord-346267-l08ld2cy 198 37 + + CC cord-346267-l08ld2cy 198 38 Γ γ NN cord-346267-l08ld2cy 198 39 4 4 CD cord-346267-l08ld2cy 198 40 ( ( -LRB- cord-346267-l08ld2cy 198 41 fig fig NN cord-346267-l08ld2cy 198 42 . . . cord-346267-l08ld2cy 198 43 3 3 CD cord-346267-l08ld2cy 198 44 ) ) -RRB- cord-346267-l08ld2cy 198 45 . . . cord-346267-l08ld2cy 199 1 Importantly importantly RB cord-346267-l08ld2cy 199 2 , , , cord-346267-l08ld2cy 199 3 the the DT cord-346267-l08ld2cy 199 4 lengths length NNS cord-346267-l08ld2cy 199 5 of of IN cord-346267-l08ld2cy 199 6 the the DT cord-346267-l08ld2cy 199 7 more more RBR cord-346267-l08ld2cy 199 8 recent recent JJ cord-346267-l08ld2cy 199 9 intraspecies intraspecie NNS cord-346267-l08ld2cy 199 10 / / SYM cord-346267-l08ld2cy 199 11 subtype subtype NN cord-346267-l08ld2cy 199 12 / / SYM cord-346267-l08ld2cy 199 13 serotype serotype NN cord-346267-l08ld2cy 199 14 branches branch NNS cord-346267-l08ld2cy 199 15 were be VBD cord-346267-l08ld2cy 199 16 relatively relatively RB cord-346267-l08ld2cy 199 17 unchanged unchanged JJ cord-346267-l08ld2cy 199 18 between between IN cord-346267-l08ld2cy 199 19 these these DT cord-346267-l08ld2cy 199 20 two two CD cord-346267-l08ld2cy 199 21 models model NNS cord-346267-l08ld2cy 199 22 for for IN cord-346267-l08ld2cy 199 23 all all DT cord-346267-l08ld2cy 199 24 three three CD cord-346267-l08ld2cy 199 25 data datum NNS cord-346267-l08ld2cy 199 26 sets set NNS cord-346267-l08ld2cy 199 27 ; ; : cord-346267-l08ld2cy 199 28 this this DT cord-346267-l08ld2cy 199 29 observation observation NN cord-346267-l08ld2cy 199 30 confirms confirm VBZ cord-346267-l08ld2cy 199 31 that that IN cord-346267-l08ld2cy 199 32 when when WRB cord-346267-l08ld2cy 199 33 the the DT cord-346267-l08ld2cy 199 34 phylogeny phylogeny NN cord-346267-l08ld2cy 199 35 is be VBZ cord-346267-l08ld2cy 199 36 comprised comprise VBN cord-346267-l08ld2cy 199 37 of of IN cord-346267-l08ld2cy 199 38 relatively relatively RB cord-346267-l08ld2cy 199 39 short short JJ cord-346267-l08ld2cy 199 40 branches branch NNS cord-346267-l08ld2cy 199 41 , , , cord-346267-l08ld2cy 199 42 it -PRON- PRP cord-346267-l08ld2cy 199 43 is be VBZ cord-346267-l08ld2cy 199 44 appropriate appropriate JJ cord-346267-l08ld2cy 199 45 to to TO cord-346267-l08ld2cy 199 46 rely rely VB cord-346267-l08ld2cy 199 47 on on IN cord-346267-l08ld2cy 199 48 common common JJ cord-346267-l08ld2cy 199 49 approximations approximation NNS cord-346267-l08ld2cy 199 50 , , , cord-346267-l08ld2cy 199 51 such such JJ cord-346267-l08ld2cy 199 52 as as IN cord-346267-l08ld2cy 199 53 the the DT cord-346267-l08ld2cy 199 54 GTR GTR NNP cord-346267-l08ld2cy 199 55 + + SYM cord-346267-l08ld2cy 199 56 Γ γ NN cord-346267-l08ld2cy 199 57 4 4 CD cord-346267-l08ld2cy 199 58 model model NN cord-346267-l08ld2cy 199 59 . . . cord-346267-l08ld2cy 200 1 This this DT cord-346267-l08ld2cy 200 2 increase increase NN cord-346267-l08ld2cy 200 3 in in IN cord-346267-l08ld2cy 200 4 the the DT cord-346267-l08ld2cy 200 5 length length NN cord-346267-l08ld2cy 200 6 of of IN cord-346267-l08ld2cy 200 7 only only JJ cord-346267-l08ld2cy 200 8 deep deep JJ cord-346267-l08ld2cy 200 9 internal internal JJ cord-346267-l08ld2cy 200 10 branches branch NNS cord-346267-l08ld2cy 200 11 was be VBD cord-346267-l08ld2cy 200 12 likely likely JJ cord-346267-l08ld2cy 200 13 due due IN cord-346267-l08ld2cy 200 14 to to IN cord-346267-l08ld2cy 200 15 the the DT cord-346267-l08ld2cy 200 16 dramatically dramatically RB cord-346267-l08ld2cy 200 17 stronger strong JJR cord-346267-l08ld2cy 200 18 purifying purifying NN cord-346267-l08ld2cy 200 19 selection selection NN cord-346267-l08ld2cy 200 20 which which WDT cord-346267-l08ld2cy 200 21 we -PRON- PRP cord-346267-l08ld2cy 200 22 inferred infer VBD cord-346267-l08ld2cy 200 23 along along IN cord-346267-l08ld2cy 200 24 these these DT cord-346267-l08ld2cy 200 25 lineages lineage NNS cord-346267-l08ld2cy 200 26 ( ( -LRB- cord-346267-l08ld2cy 200 27 table table NN cord-346267-l08ld2cy 200 28 2 2 CD cord-346267-l08ld2cy 200 29 ) ) -RRB- cord-346267-l08ld2cy 200 30 . . . cord-346267-l08ld2cy 201 1 In in IN cord-346267-l08ld2cy 201 2 AIV AIV NNP cord-346267-l08ld2cy 201 3 , , , cord-346267-l08ld2cy 201 4 for for IN cord-346267-l08ld2cy 201 5 instance instance NN cord-346267-l08ld2cy 201 6 , , , cord-346267-l08ld2cy 201 7 E e LS cord-346267-l08ld2cy 201 8 [ [ -LRB- cord-346267-l08ld2cy 201 9 β]/E β]/e XX cord-346267-l08ld2cy 201 10 [ [ -LRB- cord-346267-l08ld2cy 201 11 α α NN cord-346267-l08ld2cy 201 12 ] ] -RRB- cord-346267-l08ld2cy 201 13 was be VBD cord-346267-l08ld2cy 201 14 two two CD cord-346267-l08ld2cy 201 15 orders order NNS cord-346267-l08ld2cy 201 16 of of IN cord-346267-l08ld2cy 201 17 magnitude magnitude NN cord-346267-l08ld2cy 201 18 lower low JJR cord-346267-l08ld2cy 201 19 on on IN cord-346267-l08ld2cy 201 20 deep deep JJ cord-346267-l08ld2cy 201 21 interior interior JJ cord-346267-l08ld2cy 201 22 branches branch NNS cord-346267-l08ld2cy 201 23 , , , cord-346267-l08ld2cy 201 24 compared compare VBN cord-346267-l08ld2cy 201 25 with with IN cord-346267-l08ld2cy 201 26 the the DT cord-346267-l08ld2cy 201 27 rest rest NN cord-346267-l08ld2cy 201 28 of of IN cord-346267-l08ld2cy 201 29 the the DT cord-346267-l08ld2cy 201 30 tree tree NN cord-346267-l08ld2cy 201 31 . . . cord-346267-l08ld2cy 202 1 One one CD cord-346267-l08ld2cy 202 2 outcome outcome NN cord-346267-l08ld2cy 202 3 of of IN cord-346267-l08ld2cy 202 4 including include VBG cord-346267-l08ld2cy 202 5 variation variation NN cord-346267-l08ld2cy 202 6 in in IN cord-346267-l08ld2cy 202 7 selection selection NN cord-346267-l08ld2cy 202 8 pressures pressure NNS cord-346267-l08ld2cy 202 9 across across IN cord-346267-l08ld2cy 202 10 branches branch NNS cord-346267-l08ld2cy 202 11 into into IN cord-346267-l08ld2cy 202 12 the the DT cord-346267-l08ld2cy 202 13 evolutionary evolutionary JJ cord-346267-l08ld2cy 202 14 model model NN cord-346267-l08ld2cy 202 15 was be VBD cord-346267-l08ld2cy 202 16 that that IN cord-346267-l08ld2cy 202 17 several several JJ cord-346267-l08ld2cy 202 18 branches branch NNS cord-346267-l08ld2cy 202 19 were be VBD cord-346267-l08ld2cy 202 20 inferred infer VBN cord-346267-l08ld2cy 202 21 to to TO cord-346267-l08ld2cy 202 22 have have VB cord-346267-l08ld2cy 202 23 essentially essentially RB cord-346267-l08ld2cy 202 24 infinite infinite JJ cord-346267-l08ld2cy 202 25 lengths length NNS cord-346267-l08ld2cy 202 26 in in IN cord-346267-l08ld2cy 202 27 the the DT cord-346267-l08ld2cy 202 28 EBOV EBOV NNP cord-346267-l08ld2cy 202 29 and and CC cord-346267-l08ld2cy 202 30 AIV AIV NNP cord-346267-l08ld2cy 202 31 trees tree NNS cord-346267-l08ld2cy 202 32 . . . cord-346267-l08ld2cy 203 1 The the DT cord-346267-l08ld2cy 203 2 inference inference NN cord-346267-l08ld2cy 203 3 of of IN cord-346267-l08ld2cy 203 4 an an DT cord-346267-l08ld2cy 203 5 infinite infinite JJ cord-346267-l08ld2cy 203 6 branch branch NN cord-346267-l08ld2cy 203 7 length length NN cord-346267-l08ld2cy 203 8 is be VBZ cord-346267-l08ld2cy 203 9 likely likely RB cord-346267-l08ld2cy 203 10 caused cause VBN cord-346267-l08ld2cy 203 11 by by IN cord-346267-l08ld2cy 203 12 the the DT cord-346267-l08ld2cy 203 13 complete complete JJ cord-346267-l08ld2cy 203 14 saturation saturation NN cord-346267-l08ld2cy 203 15 of of IN cord-346267-l08ld2cy 203 16 synonymous synonymous JJ cord-346267-l08ld2cy 203 17 substitutions substitution NNS cord-346267-l08ld2cy 203 18 along along IN cord-346267-l08ld2cy 203 19 the the DT cord-346267-l08ld2cy 203 20 branch branch NN cord-346267-l08ld2cy 203 21 in in IN cord-346267-l08ld2cy 203 22 question question NN cord-346267-l08ld2cy 203 23 ; ; : cord-346267-l08ld2cy 203 24 even even RB cord-346267-l08ld2cy 203 25 after after IN cord-346267-l08ld2cy 203 26 accounting account VBG cord-346267-l08ld2cy 203 27 for for IN cord-346267-l08ld2cy 203 28 variation variation NN cord-346267-l08ld2cy 203 29 in in IN cord-346267-l08ld2cy 203 30 α α NNP cord-346267-l08ld2cy 203 31 s s NNP cord-346267-l08ld2cy 203 32 and and CC cord-346267-l08ld2cy 203 33 β β NNP cord-346267-l08ld2cy 203 34 s s NNP cord-346267-l08ld2cy 203 35 , , , cord-346267-l08ld2cy 203 36 the the DT cord-346267-l08ld2cy 203 37 true true JJ cord-346267-l08ld2cy 203 38 branch branch NN cord-346267-l08ld2cy 203 39 length length NN cord-346267-l08ld2cy 203 40 was be VBD cord-346267-l08ld2cy 203 41 inestimable inestimable JJ cord-346267-l08ld2cy 203 42 . . . cord-346267-l08ld2cy 204 1 In in IN cord-346267-l08ld2cy 204 2 the the DT cord-346267-l08ld2cy 204 3 EBOV EBOV NNP cord-346267-l08ld2cy 204 4 phylogeny phylogeny NN cord-346267-l08ld2cy 204 5 , , , cord-346267-l08ld2cy 204 6 complete complete JJ cord-346267-l08ld2cy 204 7 saturation saturation NN cord-346267-l08ld2cy 204 8 was be VBD cord-346267-l08ld2cy 204 9 observed observe VBN cord-346267-l08ld2cy 204 10 on on IN cord-346267-l08ld2cy 204 11 the the DT cord-346267-l08ld2cy 204 12 branch branch NN cord-346267-l08ld2cy 204 13 leading lead VBG cord-346267-l08ld2cy 204 14 to to IN cord-346267-l08ld2cy 204 15 EBOV EBOV NNP cord-346267-l08ld2cy 204 16 Sudan Sudan NNP cord-346267-l08ld2cy 204 17 and and CC cord-346267-l08ld2cy 204 18 on on IN cord-346267-l08ld2cy 204 19 the the DT cord-346267-l08ld2cy 204 20 branch branch NN cord-346267-l08ld2cy 204 21 connecting connect VBG cord-346267-l08ld2cy 204 22 EBOV EBOV NNP cord-346267-l08ld2cy 204 23 Reston Reston NNP cord-346267-l08ld2cy 204 24 / / SYM cord-346267-l08ld2cy 204 25 Sudan Sudan NNP cord-346267-l08ld2cy 204 26 to to IN cord-346267-l08ld2cy 204 27 EBOV EBOV NNP cord-346267-l08ld2cy 204 28 Zaire Zaire NNP cord-346267-l08ld2cy 204 29 / / SYM cord-346267-l08ld2cy 204 30 Côte Côte NNP cord-346267-l08ld2cy 204 31 d'Ivoire d'Ivoire NNP cord-346267-l08ld2cy 204 32 / / SYM cord-346267-l08ld2cy 204 33 Bundibugyo Bundibugyo NNP cord-346267-l08ld2cy 204 34 ( ( -LRB- cord-346267-l08ld2cy 204 35 supplementary supplementary JJ cord-346267-l08ld2cy 204 36 fig fig NN cord-346267-l08ld2cy 204 37 . . . cord-346267-l08ld2cy 205 1 S2 S2 NNP cord-346267-l08ld2cy 205 2 , , , cord-346267-l08ld2cy 205 3 Supplementary Supplementary NNP cord-346267-l08ld2cy 205 4 Material Material NNP cord-346267-l08ld2cy 205 5 online online RB cord-346267-l08ld2cy 205 6 ) ) -RRB- cord-346267-l08ld2cy 205 7 . . . cord-346267-l08ld2cy 206 1 In in IN cord-346267-l08ld2cy 206 2 the the DT cord-346267-l08ld2cy 206 3 AIV AIV NNP cord-346267-l08ld2cy 206 4 phylogeny phylogeny NN cord-346267-l08ld2cy 206 5 , , , cord-346267-l08ld2cy 206 6 each each DT cord-346267-l08ld2cy 206 7 of of IN cord-346267-l08ld2cy 206 8 the the DT cord-346267-l08ld2cy 206 9 nine nine CD cord-346267-l08ld2cy 206 10 neuraminidase neuraminidase NN cord-346267-l08ld2cy 206 11 serotypes serotype NNS cord-346267-l08ld2cy 206 12 naturally naturally RB cord-346267-l08ld2cy 206 13 found find VBN cord-346267-l08ld2cy 206 14 in in IN cord-346267-l08ld2cy 206 15 avian avian JJ cord-346267-l08ld2cy 206 16 hosts host NNS cord-346267-l08ld2cy 206 17 was be VBD cord-346267-l08ld2cy 206 18 separated separate VBN cord-346267-l08ld2cy 206 19 by by IN cord-346267-l08ld2cy 206 20 branches branch NNS cord-346267-l08ld2cy 206 21 that that WDT cord-346267-l08ld2cy 206 22 experienced experience VBD cord-346267-l08ld2cy 206 23 saturation saturation NN cord-346267-l08ld2cy 206 24 at at IN cord-346267-l08ld2cy 206 25 synonymous synonymous JJ cord-346267-l08ld2cy 206 26 sites site NNS cord-346267-l08ld2cy 206 27 under under IN cord-346267-l08ld2cy 206 28 the the DT cord-346267-l08ld2cy 206 29 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 206 30 model model NN cord-346267-l08ld2cy 206 31 ( ( -LRB- cord-346267-l08ld2cy 206 32 supplementary supplementary JJ cord-346267-l08ld2cy 206 33 fig fig NN cord-346267-l08ld2cy 206 34 . . . cord-346267-l08ld2cy 207 1 S3 S3 NNP cord-346267-l08ld2cy 207 2 , , , cord-346267-l08ld2cy 207 3 Supplementary Supplementary NNP cord-346267-l08ld2cy 207 4 Material Material NNP cord-346267-l08ld2cy 207 5 online online RB cord-346267-l08ld2cy 207 6 ) ) -RRB- cord-346267-l08ld2cy 207 7 . . . cord-346267-l08ld2cy 208 1 The the DT cord-346267-l08ld2cy 208 2 Latin latin JJ cord-346267-l08ld2cy 208 3 hypercube hypercube NN cord-346267-l08ld2cy 208 4 resampling resampling NN cord-346267-l08ld2cy 208 5 scheme scheme NN cord-346267-l08ld2cy 208 6 suggested suggest VBD cord-346267-l08ld2cy 208 7 relatively relatively RB cord-346267-l08ld2cy 208 8 narrow narrow JJ cord-346267-l08ld2cy 208 9 variance variance NN cord-346267-l08ld2cy 208 10 in in IN cord-346267-l08ld2cy 208 11 the the DT cord-346267-l08ld2cy 208 12 expansion expansion NN cord-346267-l08ld2cy 208 13 under under IN cord-346267-l08ld2cy 208 14 the the DT cord-346267-l08ld2cy 208 15 Lin Lin NNP cord-346267-l08ld2cy 208 16 - - HYPH cord-346267-l08ld2cy 208 17 eage+Dual eage+Dual NNP cord-346267-l08ld2cy 208 18 model model NN cord-346267-l08ld2cy 208 19 . . . cord-346267-l08ld2cy 209 1 The the DT cord-346267-l08ld2cy 209 2 total total JJ cord-346267-l08ld2cy 209 3 increase increase NN cord-346267-l08ld2cy 209 4 in in IN cord-346267-l08ld2cy 209 5 MeV MeV NNP cord-346267-l08ld2cy 209 6 / / SYM cord-346267-l08ld2cy 209 7 RPV RPV NNP cord-346267-l08ld2cy 209 8 / / SYM cord-346267-l08ld2cy 209 9 PPRV PPRV NNP cord-346267-l08ld2cy 209 10 tree tree NN cord-346267-l08ld2cy 209 11 relative relative JJ cord-346267-l08ld2cy 209 12 to to IN cord-346267-l08ld2cy 209 13 GTR GTR NNP cord-346267-l08ld2cy 209 14 + + CC cord-346267-l08ld2cy 209 15 Γ γ NN cord-346267-l08ld2cy 209 16 4 4 CD cord-346267-l08ld2cy 209 17 was be VBD cord-346267-l08ld2cy 209 18 1.93 1.93 CD cord-346267-l08ld2cy 209 19 ( ( -LRB- cord-346267-l08ld2cy 209 20 approximate approximate JJ cord-346267-l08ld2cy 209 21 95 95 CD cord-346267-l08ld2cy 209 22 % % NN cord-346267-l08ld2cy 209 23 confidence confidence NN cord-346267-l08ld2cy 209 24 interval interval NN cord-346267-l08ld2cy 209 25 : : : cord-346267-l08ld2cy 209 26 1.76 1.76 CD cord-346267-l08ld2cy 209 27 - - HYPH cord-346267-l08ld2cy 209 28 2.16 2.16 CD cord-346267-l08ld2cy 209 29 ) ) -RRB- cord-346267-l08ld2cy 209 30 . . . cord-346267-l08ld2cy 210 1 Due due IN cord-346267-l08ld2cy 210 2 to to IN cord-346267-l08ld2cy 210 3 the the DT cord-346267-l08ld2cy 210 4 inference inference NN cord-346267-l08ld2cy 210 5 of of IN cord-346267-l08ld2cy 210 6 branch branch NN cord-346267-l08ld2cy 210 7 lengths length NNS cord-346267-l08ld2cy 210 8 that that WDT cord-346267-l08ld2cy 210 9 experienced experience VBD cord-346267-l08ld2cy 210 10 saturation saturation NN cord-346267-l08ld2cy 210 11 in in IN cord-346267-l08ld2cy 210 12 EBOV EBOV NNP cord-346267-l08ld2cy 210 13 and and CC cord-346267-l08ld2cy 210 14 AIV AIV NNP cord-346267-l08ld2cy 210 15 , , , cord-346267-l08ld2cy 210 16 the the DT cord-346267-l08ld2cy 210 17 overall overall JJ cord-346267-l08ld2cy 210 18 increase increase NN cord-346267-l08ld2cy 210 19 in in IN cord-346267-l08ld2cy 210 20 tree tree NN cord-346267-l08ld2cy 210 21 length length NN cord-346267-l08ld2cy 210 22 was be VBD cord-346267-l08ld2cy 210 23 more more RBR cord-346267-l08ld2cy 210 24 dramatic dramatic JJ cord-346267-l08ld2cy 210 25 under under IN cord-346267-l08ld2cy 210 26 the the DT cord-346267-l08ld2cy 210 27 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 210 28 model model NN cord-346267-l08ld2cy 210 29 , , , cord-346267-l08ld2cy 210 30 relative relative JJ cord-346267-l08ld2cy 210 31 to to IN cord-346267-l08ld2cy 210 32 GTR GTR NNP cord-346267-l08ld2cy 210 33 + + CC cord-346267-l08ld2cy 210 34 Γ γ NN cord-346267-l08ld2cy 210 35 4 4 CD cord-346267-l08ld2cy 210 36 : : : cord-346267-l08ld2cy 210 37 32.14 32.14 CD cord-346267-l08ld2cy 210 38 ( ( -LRB- cord-346267-l08ld2cy 210 39 approximate approximate JJ cord-346267-l08ld2cy 210 40 95 95 CD cord-346267-l08ld2cy 210 41 % % NN cord-346267-l08ld2cy 210 42 confidence confidence NN cord-346267-l08ld2cy 210 43 interval interval NN cord-346267-l08ld2cy 210 44 : : : cord-346267-l08ld2cy 210 45 12.14 12.14 CD cord-346267-l08ld2cy 210 46 - - SYM cord-346267-l08ld2cy 210 47 32.22 32.22 CD cord-346267-l08ld2cy 210 48 ) ) -RRB- cord-346267-l08ld2cy 210 49 for for IN cord-346267-l08ld2cy 210 50 EBOV EBOV NNP cord-346267-l08ld2cy 210 51 and and CC cord-346267-l08ld2cy 210 52 107.34 107.34 CD cord-346267-l08ld2cy 210 53 ( ( -LRB- cord-346267-l08ld2cy 210 54 approximate approximate JJ cord-346267-l08ld2cy 210 55 95 95 CD cord-346267-l08ld2cy 210 56 % % NN cord-346267-l08ld2cy 210 57 confidence confidence NN cord-346267-l08ld2cy 210 58 interval interval NN cord-346267-l08ld2cy 210 59 : : : cord-346267-l08ld2cy 210 60 101.06 101.06 CD cord-346267-l08ld2cy 210 61 - - SYM cord-346267-l08ld2cy 210 62 117.36 117.36 CD cord-346267-l08ld2cy 210 63 ) ) -RRB- cord-346267-l08ld2cy 210 64 for for IN cord-346267-l08ld2cy 210 65 AIV AIV NNP cord-346267-l08ld2cy 210 66 . . . cord-346267-l08ld2cy 211 1 It -PRON- PRP cord-346267-l08ld2cy 211 2 is be VBZ cord-346267-l08ld2cy 211 3 clear clear JJ cord-346267-l08ld2cy 211 4 that that IN cord-346267-l08ld2cy 211 5 purifying purifying NN cord-346267-l08ld2cy 211 6 selection selection NN cord-346267-l08ld2cy 211 7 can can MD cord-346267-l08ld2cy 211 8 obscure obscure VB cord-346267-l08ld2cy 211 9 the the DT cord-346267-l08ld2cy 211 10 ancient ancient JJ cord-346267-l08ld2cy 211 11 evolution evolution NN cord-346267-l08ld2cy 211 12 of of IN cord-346267-l08ld2cy 211 13 the the DT cord-346267-l08ld2cy 211 14 RNA RNA NNP cord-346267-l08ld2cy 211 15 viruses virus NNS cord-346267-l08ld2cy 211 16 examined examine VBN cord-346267-l08ld2cy 211 17 here here RB cord-346267-l08ld2cy 211 18 . . . cord-346267-l08ld2cy 212 1 A a DT cord-346267-l08ld2cy 212 2 comparison comparison NN cord-346267-l08ld2cy 212 3 of of IN cord-346267-l08ld2cy 212 4 the the DT cord-346267-l08ld2cy 212 5 MeV MeV NNP cord-346267-l08ld2cy 212 6 / / SYM cord-346267-l08ld2cy 212 7 RPV RPV NNP cord-346267-l08ld2cy 212 8 / / SYM cord-346267-l08ld2cy 212 9 PPRV PPRV NNP cord-346267-l08ld2cy 212 10 phylogeny phylogeny NN cord-346267-l08ld2cy 212 11 optimized optimize VBN cord-346267-l08ld2cy 212 12 under under IN cord-346267-l08ld2cy 212 13 GTR GTR NNP cord-346267-l08ld2cy 212 14 + + CC cord-346267-l08ld2cy 212 15 Γ γ NN cord-346267-l08ld2cy 212 16 4 4 CD cord-346267-l08ld2cy 212 17 and and CC cord-346267-l08ld2cy 212 18 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 212 19 ( ( -LRB- cord-346267-l08ld2cy 212 20 two two CD cord-346267-l08ld2cy 212 21 - - HYPH cord-346267-l08ld2cy 212 22 rate rate NN cord-346267-l08ld2cy 212 23 ) ) -RRB- cord-346267-l08ld2cy 212 24 models model NNS cord-346267-l08ld2cy 212 25 showed show VBD cord-346267-l08ld2cy 212 26 that that IN cord-346267-l08ld2cy 212 27 the the DT cord-346267-l08ld2cy 212 28 elongation elongation NN cord-346267-l08ld2cy 212 29 of of IN cord-346267-l08ld2cy 212 30 branches branch NNS cord-346267-l08ld2cy 212 31 occurred occur VBD cord-346267-l08ld2cy 212 32 along along IN cord-346267-l08ld2cy 212 33 the the DT cord-346267-l08ld2cy 212 34 deep deep JJ cord-346267-l08ld2cy 212 35 internal internal JJ cord-346267-l08ld2cy 212 36 branches branch NNS cord-346267-l08ld2cy 212 37 , , , cord-346267-l08ld2cy 212 38 leaving leave VBG cord-346267-l08ld2cy 212 39 the the DT cord-346267-l08ld2cy 212 40 relationships relationship NNS cord-346267-l08ld2cy 212 41 among among IN cord-346267-l08ld2cy 212 42 recently recently RB cord-346267-l08ld2cy 212 43 divergent divergent JJ cord-346267-l08ld2cy 212 44 lineages lineage NNS cord-346267-l08ld2cy 212 45 within within IN cord-346267-l08ld2cy 212 46 viral viral JJ cord-346267-l08ld2cy 212 47 species specie NNS cord-346267-l08ld2cy 212 48 relatively relatively RB cord-346267-l08ld2cy 212 49 unchanged unchanged JJ cord-346267-l08ld2cy 212 50 ( ( -LRB- cord-346267-l08ld2cy 212 51 fig fig NN cord-346267-l08ld2cy 212 52 . . . cord-346267-l08ld2cy 212 53 4 4 LS cord-346267-l08ld2cy 212 54 ) ) -RRB- cord-346267-l08ld2cy 212 55 . . . cord-346267-l08ld2cy 213 1 The the DT cord-346267-l08ld2cy 213 2 depth depth NN cord-346267-l08ld2cy 213 3 of of IN cord-346267-l08ld2cy 213 4 the the DT cord-346267-l08ld2cy 213 5 split split NN cord-346267-l08ld2cy 213 6 between between IN cord-346267-l08ld2cy 213 7 MeV MeV NNP cord-346267-l08ld2cy 213 8 and and CC cord-346267-l08ld2cy 213 9 RPV RPV NNP cord-346267-l08ld2cy 213 10 under under IN cord-346267-l08ld2cy 213 11 a a DT cord-346267-l08ld2cy 213 12 GTR GTR NNP cord-346267-l08ld2cy 213 13 + + SYM cord-346267-l08ld2cy 213 14 Γ γ NN cord-346267-l08ld2cy 213 15 4 4 CD cord-346267-l08ld2cy 213 16 model model NN cord-346267-l08ld2cy 213 17 was be VBD cord-346267-l08ld2cy 213 18 0.4921 0.4921 CD cord-346267-l08ld2cy 213 19 substitutions substitution NNS cord-346267-l08ld2cy 213 20 / / SYM cord-346267-l08ld2cy 213 21 site site NN cord-346267-l08ld2cy 213 22 , , , cord-346267-l08ld2cy 213 23 which which WDT cord-346267-l08ld2cy 213 24 , , , cord-346267-l08ld2cy 213 25 according accord VBG cord-346267-l08ld2cy 213 26 to to IN cord-346267-l08ld2cy 213 27 the the DT cord-346267-l08ld2cy 213 28 BMCMC BMCMC NNP cord-346267-l08ld2cy 213 29 analysis analysis NN cord-346267-l08ld2cy 213 30 , , , cord-346267-l08ld2cy 213 31 occurred occur VBD cord-346267-l08ld2cy 213 32 in in IN cord-346267-l08ld2cy 213 33 the the DT cord-346267-l08ld2cy 213 34 year year NN cord-346267-l08ld2cy 213 35 1483 1483 CD cord-346267-l08ld2cy 213 36 Table Table NNP cord-346267-l08ld2cy 213 37 2 2 CD cord-346267-l08ld2cy 213 38 . . . cord-346267-l08ld2cy 214 1 Goodness goodness NN cord-346267-l08ld2cy 214 2 of of IN cord-346267-l08ld2cy 214 3 fit fit NN cord-346267-l08ld2cy 214 4 for for IN cord-346267-l08ld2cy 214 5 various various JJ cord-346267-l08ld2cy 214 6 codon codon NN cord-346267-l08ld2cy 214 7 models model NNS cord-346267-l08ld2cy 214 8 and and CC cord-346267-l08ld2cy 214 9 the the DT cord-346267-l08ld2cy 214 10 effect effect NN cord-346267-l08ld2cy 214 11 of of IN cord-346267-l08ld2cy 214 12 model model NN cord-346267-l08ld2cy 214 13 choice choice NN cord-346267-l08ld2cy 214 14 on on IN cord-346267-l08ld2cy 214 15 the the DT cord-346267-l08ld2cy 214 16 estimates estimate NNS cord-346267-l08ld2cy 214 17 of of IN cord-346267-l08ld2cy 214 18 the the DT cord-346267-l08ld2cy 214 19 branch branch NN cord-346267-l08ld2cy 214 20 lengths length NNS cord-346267-l08ld2cy 214 21 of of IN cord-346267-l08ld2cy 214 22 deep deep JJ cord-346267-l08ld2cy 214 23 and and CC cord-346267-l08ld2cy 214 24 recent recent JJ cord-346267-l08ld2cy 214 25 lineages lineage NNS cord-346267-l08ld2cy 214 26 . . . cord-346267-l08ld2cy 215 1 Model model NN cord-346267-l08ld2cy 215 2 ( ( -LRB- cord-346267-l08ld2cy 215 3 597 597 CD cord-346267-l08ld2cy 215 4 - - SYM cord-346267-l08ld2cy 215 5 1144 1144 CD cord-346267-l08ld2cy 215 6 CE CE NNP cord-346267-l08ld2cy 215 7 ) ) -RRB- cord-346267-l08ld2cy 215 8 . . . cord-346267-l08ld2cy 216 1 This this DT cord-346267-l08ld2cy 216 2 extrapolation extrapolation NN cord-346267-l08ld2cy 216 3 is be VBZ cord-346267-l08ld2cy 216 4 meant mean VBN cord-346267-l08ld2cy 216 5 as as IN cord-346267-l08ld2cy 216 6 a a DT cord-346267-l08ld2cy 216 7 rather rather RB cord-346267-l08ld2cy 216 8 crude crude JJ cord-346267-l08ld2cy 216 9 approximation approximation NN cord-346267-l08ld2cy 216 10 of of IN cord-346267-l08ld2cy 216 11 the the DT cord-346267-l08ld2cy 216 12 age age NN cord-346267-l08ld2cy 216 13 of of IN cord-346267-l08ld2cy 216 14 this this DT cord-346267-l08ld2cy 216 15 divergence divergence NN cord-346267-l08ld2cy 216 16 event event NN cord-346267-l08ld2cy 216 17 ; ; : cord-346267-l08ld2cy 216 18 nevertheless nevertheless RB cord-346267-l08ld2cy 216 19 , , , cord-346267-l08ld2cy 216 20 these these DT cord-346267-l08ld2cy 216 21 dates date NNS cord-346267-l08ld2cy 216 22 are be VBP cord-346267-l08ld2cy 216 23 more more RBR cord-346267-l08ld2cy 216 24 likely likely JJ cord-346267-l08ld2cy 216 25 to to TO cord-346267-l08ld2cy 216 26 be be VB cord-346267-l08ld2cy 216 27 closer close JJR cord-346267-l08ld2cy 216 28 to to IN cord-346267-l08ld2cy 216 29 the the DT cord-346267-l08ld2cy 216 30 true true JJ cord-346267-l08ld2cy 216 31 split split NN cord-346267-l08ld2cy 216 32 between between IN cord-346267-l08ld2cy 216 33 MeV MeV NNP cord-346267-l08ld2cy 216 34 and and CC cord-346267-l08ld2cy 216 35 RPV RPV NNP cord-346267-l08ld2cy 216 36 than than IN cord-346267-l08ld2cy 216 37 previous previous JJ cord-346267-l08ld2cy 216 38 estimates estimate NNS cord-346267-l08ld2cy 216 39 and and CC cord-346267-l08ld2cy 216 40 are be VBP cord-346267-l08ld2cy 216 41 not not RB cord-346267-l08ld2cy 216 42 inconsistent inconsistent JJ cord-346267-l08ld2cy 216 43 with with IN cord-346267-l08ld2cy 216 44 recorded record VBN cord-346267-l08ld2cy 216 45 history history NN cord-346267-l08ld2cy 216 46 of of IN cord-346267-l08ld2cy 216 47 measles measle NNS cord-346267-l08ld2cy 216 48 . . . cord-346267-l08ld2cy 217 1 The the DT cord-346267-l08ld2cy 217 2 degree degree NN cord-346267-l08ld2cy 217 3 of of IN cord-346267-l08ld2cy 217 4 synonymous synonymous JJ cord-346267-l08ld2cy 217 5 saturation saturation NN cord-346267-l08ld2cy 217 6 observed observe VBN cord-346267-l08ld2cy 217 7 along along IN cord-346267-l08ld2cy 217 8 the the DT cord-346267-l08ld2cy 217 9 EBOV EBOV NNP cord-346267-l08ld2cy 217 10 and and CC cord-346267-l08ld2cy 217 11 AIV AIV NNP cord-346267-l08ld2cy 217 12 phylogenies phylogeny NNS cord-346267-l08ld2cy 217 13 indicate indicate VBP cord-346267-l08ld2cy 217 14 an an DT cord-346267-l08ld2cy 217 15 inability inability NN cord-346267-l08ld2cy 217 16 to to TO cord-346267-l08ld2cy 217 17 reliably reliably RB cord-346267-l08ld2cy 217 18 infer infer VB cord-346267-l08ld2cy 217 19 tMRCAs tmrcas CD cord-346267-l08ld2cy 217 20 for for IN cord-346267-l08ld2cy 217 21 these these DT cord-346267-l08ld2cy 217 22 viruses virus NNS cord-346267-l08ld2cy 217 23 . . . cord-346267-l08ld2cy 218 1 Too too RB cord-346267-l08ld2cy 218 2 many many JJ cord-346267-l08ld2cy 218 3 sites site NNS cord-346267-l08ld2cy 218 4 have have VBP cord-346267-l08ld2cy 218 5 sunk sink VBN cord-346267-l08ld2cy 218 6 beyond beyond IN cord-346267-l08ld2cy 218 7 the the DT cord-346267-l08ld2cy 218 8 evolutionary evolutionary JJ cord-346267-l08ld2cy 218 9 horizon horizon NN cord-346267-l08ld2cy 218 10 . . . cord-346267-l08ld2cy 219 1 Nevertheless nevertheless RB cord-346267-l08ld2cy 219 2 , , , cord-346267-l08ld2cy 219 3 these these DT cord-346267-l08ld2cy 219 4 estimates estimate NNS cord-346267-l08ld2cy 219 5 could could MD cord-346267-l08ld2cy 219 6 provide provide VB cord-346267-l08ld2cy 219 7 meaningful meaningful JJ cord-346267-l08ld2cy 219 8 minimum minimum JJ cord-346267-l08ld2cy 219 9 bounds bound NNS cord-346267-l08ld2cy 219 10 for for IN cord-346267-l08ld2cy 219 11 the the DT cord-346267-l08ld2cy 219 12 tMRCAs tmrcas NN cord-346267-l08ld2cy 219 13 . . . cord-346267-l08ld2cy 220 1 Thus thus RB cord-346267-l08ld2cy 220 2 , , , cord-346267-l08ld2cy 220 3 for for IN cord-346267-l08ld2cy 220 4 both both CC cord-346267-l08ld2cy 220 5 EBOV EBOV NNP cord-346267-l08ld2cy 220 6 and and CC cord-346267-l08ld2cy 220 7 AIV AIV NNP cord-346267-l08ld2cy 220 8 , , , cord-346267-l08ld2cy 220 9 we -PRON- PRP cord-346267-l08ld2cy 220 10 applied apply VBD cord-346267-l08ld2cy 220 11 the the DT cord-346267-l08ld2cy 220 12 mean mean JJ cord-346267-l08ld2cy 220 13 substitution substitution NN cord-346267-l08ld2cy 220 14 rate rate NN cord-346267-l08ld2cy 220 15 inferred infer VBN cord-346267-l08ld2cy 220 16 in in IN cord-346267-l08ld2cy 220 17 the the DT cord-346267-l08ld2cy 220 18 BMCMC BMCMC NNP cord-346267-l08ld2cy 220 19 GTR GTR NNP cord-346267-l08ld2cy 220 20 + + CC cord-346267-l08ld2cy 220 21 Γ γ NN cord-346267-l08ld2cy 220 22 4 4 CD cord-346267-l08ld2cy 220 23 analyses analysis NNS cord-346267-l08ld2cy 220 24 to to IN cord-346267-l08ld2cy 220 25 the the DT cord-346267-l08ld2cy 220 26 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 220 27 phylogenies phylogeny NNS cord-346267-l08ld2cy 220 28 . . . cord-346267-l08ld2cy 221 1 This this DT cord-346267-l08ld2cy 221 2 approach approach NN cord-346267-l08ld2cy 221 3 suggested suggest VBD cord-346267-l08ld2cy 221 4 that that IN cord-346267-l08ld2cy 221 5 the the DT cord-346267-l08ld2cy 221 6 minimum minimum JJ cord-346267-l08ld2cy 221 7 tMRCA tmrca NN cord-346267-l08ld2cy 221 8 estimates estimate NNS cord-346267-l08ld2cy 221 9 for for IN cord-346267-l08ld2cy 221 10 EBOV EBOV NNP cord-346267-l08ld2cy 221 11 and and CC cord-346267-l08ld2cy 221 12 AIV AIV NNP cord-346267-l08ld2cy 221 13 are be VBP cord-346267-l08ld2cy 221 14 approximately approximately RB cord-346267-l08ld2cy 221 15 70,000 70,000 CD cord-346267-l08ld2cy 221 16 and and CC cord-346267-l08ld2cy 221 17 200,000 200,000 CD cord-346267-l08ld2cy 221 18 years year NNS cord-346267-l08ld2cy 221 19 ago ago RB cord-346267-l08ld2cy 221 20 , , , cord-346267-l08ld2cy 221 21 respectively respectively RB cord-346267-l08ld2cy 221 22 . . . cord-346267-l08ld2cy 222 1 The the DT cord-346267-l08ld2cy 222 2 actual actual JJ cord-346267-l08ld2cy 222 3 tMRCAs tmrcas NN cord-346267-l08ld2cy 222 4 may may MD cord-346267-l08ld2cy 222 5 be be VB cord-346267-l08ld2cy 222 6 much much RB cord-346267-l08ld2cy 222 7 older old JJR cord-346267-l08ld2cy 222 8 . . . cord-346267-l08ld2cy 223 1 Our -PRON- PRP$ cord-346267-l08ld2cy 223 2 results result NNS cord-346267-l08ld2cy 223 3 suggest suggest VBP cord-346267-l08ld2cy 223 4 that that IN cord-346267-l08ld2cy 223 5 the the DT cord-346267-l08ld2cy 223 6 ancient ancient JJ cord-346267-l08ld2cy 223 7 age age NN cord-346267-l08ld2cy 223 8 of of IN cord-346267-l08ld2cy 223 9 RNA RNA NNP cord-346267-l08ld2cy 223 10 viruses virus NNS cord-346267-l08ld2cy 223 11 may may MD cord-346267-l08ld2cy 223 12 be be VB cord-346267-l08ld2cy 223 13 partially partially RB cord-346267-l08ld2cy 223 14 masked mask VBN cord-346267-l08ld2cy 223 15 behind behind IN cord-346267-l08ld2cy 223 16 a a DT cord-346267-l08ld2cy 223 17 veil veil NN cord-346267-l08ld2cy 223 18 of of IN cord-346267-l08ld2cy 223 19 purifying purify VBG cord-346267-l08ld2cy 223 20 selection selection NN cord-346267-l08ld2cy 223 21 . . . cord-346267-l08ld2cy 224 1 The the DT cord-346267-l08ld2cy 224 2 same same JJ cord-346267-l08ld2cy 224 3 forces force NNS cord-346267-l08ld2cy 224 4 of of IN cord-346267-l08ld2cy 224 5 purifying purify VBG cord-346267-l08ld2cy 224 6 selection selection NN cord-346267-l08ld2cy 224 7 that that WDT cord-346267-l08ld2cy 224 8 maintain maintain VBP cord-346267-l08ld2cy 224 9 evidence evidence NN cord-346267-l08ld2cy 224 10 of of IN cord-346267-l08ld2cy 224 11 protein protein NN cord-346267-l08ld2cy 224 12 sequence sequence NN cord-346267-l08ld2cy 224 13 homology homology NN cord-346267-l08ld2cy 224 14 over over IN cord-346267-l08ld2cy 224 15 great great JJ cord-346267-l08ld2cy 224 16 evolutionary evolutionary JJ cord-346267-l08ld2cy 224 17 distances distance NNS cord-346267-l08ld2cy 224 18 also also RB cord-346267-l08ld2cy 224 19 truncate truncate VBP cord-346267-l08ld2cy 224 20 long long JJ cord-346267-l08ld2cy 224 21 ancestral ancestral JJ cord-346267-l08ld2cy 224 22 branches branch NNS cord-346267-l08ld2cy 224 23 deep deep RB cord-346267-l08ld2cy 224 24 within within IN cord-346267-l08ld2cy 224 25 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 224 26 trees tree NNS cord-346267-l08ld2cy 224 27 . . . cord-346267-l08ld2cy 225 1 We -PRON- PRP cord-346267-l08ld2cy 225 2 observed observe VBD cord-346267-l08ld2cy 225 3 this this DT cord-346267-l08ld2cy 225 4 pattern pattern NN cord-346267-l08ld2cy 225 5 MBE MBE NNP cord-346267-l08ld2cy 225 6 in in IN cord-346267-l08ld2cy 225 7 three three CD cord-346267-l08ld2cy 225 8 different different JJ cord-346267-l08ld2cy 225 9 groups group NNS cord-346267-l08ld2cy 225 10 of of IN cord-346267-l08ld2cy 225 11 rapidly rapidly RB cord-346267-l08ld2cy 225 12 evolving evolve VBG cord-346267-l08ld2cy 225 13 negative negative JJ cord-346267-l08ld2cy 225 14 - - HYPH cord-346267-l08ld2cy 225 15 sense sense NN cord-346267-l08ld2cy 225 16 RNA RNA NNP cord-346267-l08ld2cy 225 17 viruses virus NNS cord-346267-l08ld2cy 225 18 : : : cord-346267-l08ld2cy 226 1 MeV MeV NNP cord-346267-l08ld2cy 226 2 / / SYM cord-346267-l08ld2cy 226 3 RPV RPV NNP cord-346267-l08ld2cy 226 4 / / SYM cord-346267-l08ld2cy 226 5 PPRV PPRV NNP cord-346267-l08ld2cy 226 6 , , , cord-346267-l08ld2cy 226 7 EBOV EBOV NNP cord-346267-l08ld2cy 226 8 , , , cord-346267-l08ld2cy 226 9 and and CC cord-346267-l08ld2cy 226 10 AIV AIV NNP cord-346267-l08ld2cy 226 11 . . . cord-346267-l08ld2cy 227 1 Estimating estimate VBG cord-346267-l08ld2cy 227 2 branch branch NN cord-346267-l08ld2cy 227 3 lengths length NNS cord-346267-l08ld2cy 227 4 under under IN cord-346267-l08ld2cy 227 5 a a DT cord-346267-l08ld2cy 227 6 codon codon NN cord-346267-l08ld2cy 227 7 - - HYPH cord-346267-l08ld2cy 227 8 based base VBN cord-346267-l08ld2cy 227 9 substitution substitution NN cord-346267-l08ld2cy 227 10 model model NN cord-346267-l08ld2cy 227 11 that that WDT cord-346267-l08ld2cy 227 12 accounts account VBZ cord-346267-l08ld2cy 227 13 for for IN cord-346267-l08ld2cy 227 14 spatial spatial JJ cord-346267-l08ld2cy 227 15 and and CC cord-346267-l08ld2cy 227 16 temporal temporal JJ cord-346267-l08ld2cy 227 17 variation variation NN cord-346267-l08ld2cy 227 18 in in IN cord-346267-l08ld2cy 227 19 selection selection NN cord-346267-l08ld2cy 227 20 pressures pressure NNS cord-346267-l08ld2cy 227 21 yielded yield VBD cord-346267-l08ld2cy 227 22 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 227 23 trees tree NNS cord-346267-l08ld2cy 227 24 that that WDT cord-346267-l08ld2cy 227 25 were be VBD cord-346267-l08ld2cy 227 26 more more JJR cord-346267-l08ld2cy 227 27 than than IN cord-346267-l08ld2cy 227 28 twice twice RB cord-346267-l08ld2cy 227 29 as as RB cord-346267-l08ld2cy 227 30 long long RB cord-346267-l08ld2cy 227 31 as as IN cord-346267-l08ld2cy 227 32 those those DT cord-346267-l08ld2cy 227 33 obtained obtain VBN cord-346267-l08ld2cy 227 34 under under IN cord-346267-l08ld2cy 227 35 standard standard JJ cord-346267-l08ld2cy 227 36 nucleotide nucleotide JJ cord-346267-l08ld2cy 227 37 models model NNS cord-346267-l08ld2cy 227 38 . . . cord-346267-l08ld2cy 228 1 Modeling model VBG cord-346267-l08ld2cy 228 2 synonymous synonymous JJ cord-346267-l08ld2cy 228 3 , , , cord-346267-l08ld2cy 228 4 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 228 5 , , , cord-346267-l08ld2cy 228 6 and and CC cord-346267-l08ld2cy 228 7 lineage lineage NN cord-346267-l08ld2cy 228 8 - - HYPH cord-346267-l08ld2cy 228 9 specific specific JJ cord-346267-l08ld2cy 228 10 rate rate NN cord-346267-l08ld2cy 228 11 variation variation NN cord-346267-l08ld2cy 228 12 indicates indicate VBZ cord-346267-l08ld2cy 228 13 that that IN cord-346267-l08ld2cy 228 14 current current JJ cord-346267-l08ld2cy 228 15 estimates estimate NNS cord-346267-l08ld2cy 228 16 of of IN cord-346267-l08ld2cy 228 17 the the DT cord-346267-l08ld2cy 228 18 age age NN cord-346267-l08ld2cy 228 19 of of IN cord-346267-l08ld2cy 228 20 these these DT cord-346267-l08ld2cy 228 21 , , , cord-346267-l08ld2cy 228 22 and and CC cord-346267-l08ld2cy 228 23 possibly possibly RB cord-346267-l08ld2cy 228 24 other other JJ cord-346267-l08ld2cy 228 25 , , , cord-346267-l08ld2cy 228 26 viral viral JJ cord-346267-l08ld2cy 228 27 lineages lineage NNS cord-346267-l08ld2cy 228 28 may may MD cord-346267-l08ld2cy 228 29 be be VB cord-346267-l08ld2cy 228 30 dramatically dramatically RB cord-346267-l08ld2cy 228 31 underestimated underestimate VBN cord-346267-l08ld2cy 228 32 . . . cord-346267-l08ld2cy 229 1 Codon Codon NNP cord-346267-l08ld2cy 229 2 models model NNS cord-346267-l08ld2cy 229 3 used use VBN cord-346267-l08ld2cy 229 4 in in IN cord-346267-l08ld2cy 229 5 this this DT cord-346267-l08ld2cy 229 6 study study NN cord-346267-l08ld2cy 229 7 do do VBP cord-346267-l08ld2cy 229 8 not not RB cord-346267-l08ld2cy 229 9 circumvent circumvent VB cord-346267-l08ld2cy 229 10 the the DT cord-346267-l08ld2cy 229 11 issue issue NN cord-346267-l08ld2cy 229 12 of of IN cord-346267-l08ld2cy 229 13 substitutional substitutional JJ cord-346267-l08ld2cy 229 14 saturation saturation NN cord-346267-l08ld2cy 229 15 but but CC cord-346267-l08ld2cy 229 16 merely merely RB cord-346267-l08ld2cy 229 17 extend extend VB cord-346267-l08ld2cy 229 18 the the DT cord-346267-l08ld2cy 229 19 horizon horizon NN cord-346267-l08ld2cy 229 20 further further RB cord-346267-l08ld2cy 229 21 back back RB cord-346267-l08ld2cy 229 22 in in IN cord-346267-l08ld2cy 229 23 evolutionary evolutionary JJ cord-346267-l08ld2cy 229 24 time time NN cord-346267-l08ld2cy 229 25 ; ; : cord-346267-l08ld2cy 229 26 the the DT cord-346267-l08ld2cy 229 27 application application NN cord-346267-l08ld2cy 229 28 of of IN cord-346267-l08ld2cy 229 29 more more RBR cord-346267-l08ld2cy 229 30 complex complex JJ cord-346267-l08ld2cy 229 31 and and CC cord-346267-l08ld2cy 229 32 biologically biologically RB cord-346267-l08ld2cy 229 33 realistic realistic JJ cord-346267-l08ld2cy 229 34 models model NNS cord-346267-l08ld2cy 229 35 is be VBZ cord-346267-l08ld2cy 229 36 likely likely JJ cord-346267-l08ld2cy 229 37 to to TO cord-346267-l08ld2cy 229 38 extend extend VB cord-346267-l08ld2cy 229 39 these these DT cord-346267-l08ld2cy 229 40 branches branch NNS cord-346267-l08ld2cy 229 41 even even RB cord-346267-l08ld2cy 229 42 further far RBR cord-346267-l08ld2cy 229 43 into into IN cord-346267-l08ld2cy 229 44 the the DT cord-346267-l08ld2cy 229 45 past past NN cord-346267-l08ld2cy 229 46 . . . cord-346267-l08ld2cy 230 1 Eventually eventually RB cord-346267-l08ld2cy 230 2 , , , cord-346267-l08ld2cy 230 3 however however RB cord-346267-l08ld2cy 230 4 , , , cord-346267-l08ld2cy 230 5 even even RB cord-346267-l08ld2cy 230 6 the the DT cord-346267-l08ld2cy 230 7 most most RBS cord-346267-l08ld2cy 230 8 realistic realistic JJ cord-346267-l08ld2cy 230 9 models model NNS cord-346267-l08ld2cy 230 10 are be VBP cord-346267-l08ld2cy 230 11 likely likely JJ cord-346267-l08ld2cy 230 12 to to TO cord-346267-l08ld2cy 230 13 fail fail VB cord-346267-l08ld2cy 230 14 and and CC cord-346267-l08ld2cy 230 15 external external JJ cord-346267-l08ld2cy 230 16 information information NN cord-346267-l08ld2cy 230 17 , , , cord-346267-l08ld2cy 230 18 such such JJ cord-346267-l08ld2cy 230 19 as as IN cord-346267-l08ld2cy 230 20 biogeography biogeography NN cord-346267-l08ld2cy 230 21 and and CC cord-346267-l08ld2cy 230 22 homology homology NN cord-346267-l08ld2cy 230 23 to to IN cord-346267-l08ld2cy 230 24 endogenous endogenous JJ cord-346267-l08ld2cy 230 25 viral viral JJ cord-346267-l08ld2cy 230 26 elements element NNS cord-346267-l08ld2cy 230 27 , , , cord-346267-l08ld2cy 230 28 must must MD cord-346267-l08ld2cy 230 29 be be VB cord-346267-l08ld2cy 230 30 brought bring VBN cord-346267-l08ld2cy 230 31 into into IN cord-346267-l08ld2cy 230 32 consideration consideration NN cord-346267-l08ld2cy 230 33 to to TO cord-346267-l08ld2cy 230 34 estimate estimate VB cord-346267-l08ld2cy 230 35 truly truly RB cord-346267-l08ld2cy 230 36 ancient ancient JJ cord-346267-l08ld2cy 230 37 events event NNS cord-346267-l08ld2cy 230 38 ( ( -LRB- cord-346267-l08ld2cy 230 39 Katzourakis Katzourakis NNP cord-346267-l08ld2cy 230 40 et et NNP cord-346267-l08ld2cy 230 41 al al NNP cord-346267-l08ld2cy 230 42 . . . cord-346267-l08ld2cy 230 43 2009 2009 CD cord-346267-l08ld2cy 230 44 ; ; . cord-346267-l08ld2cy 231 1 Katzourakis Katzourakis NNP cord-346267-l08ld2cy 231 2 and and CC cord-346267-l08ld2cy 231 3 Gifford Gifford NNP cord-346267-l08ld2cy 231 4 2010 2010 CD cord-346267-l08ld2cy 231 5 ) ) -RRB- cord-346267-l08ld2cy 231 6 . . . cord-346267-l08ld2cy 232 1 The the DT cord-346267-l08ld2cy 232 2 precise precise JJ cord-346267-l08ld2cy 232 3 age age NN cord-346267-l08ld2cy 232 4 of of IN cord-346267-l08ld2cy 232 5 measles measle NNS cord-346267-l08ld2cy 232 6 in in IN cord-346267-l08ld2cy 232 7 the the DT cord-346267-l08ld2cy 232 8 human human JJ cord-346267-l08ld2cy 232 9 population population NN cord-346267-l08ld2cy 232 10 remains remain VBZ cord-346267-l08ld2cy 232 11 elusive elusive JJ cord-346267-l08ld2cy 232 12 . . . cord-346267-l08ld2cy 233 1 The the DT cord-346267-l08ld2cy 233 2 domestication domestication NN cord-346267-l08ld2cy 233 3 of of IN cord-346267-l08ld2cy 233 4 cattle cattle NNS cord-346267-l08ld2cy 233 5 beginning begin VBG cord-346267-l08ld2cy 233 6 10 10 CD cord-346267-l08ld2cy 233 7 , , , cord-346267-l08ld2cy 233 8 000 000 CD cord-346267-l08ld2cy 233 9 years year NNS cord-346267-l08ld2cy 233 10 ago ago RB cord-346267-l08ld2cy 233 11 provided provide VBD cord-346267-l08ld2cy 233 12 the the DT cord-346267-l08ld2cy 233 13 necessary necessary JJ cord-346267-l08ld2cy 233 14 exposure exposure NN cord-346267-l08ld2cy 233 15 to to IN cord-346267-l08ld2cy 233 16 RPV RPV NNP cord-346267-l08ld2cy 233 17 , , , cord-346267-l08ld2cy 233 18 and and CC cord-346267-l08ld2cy 233 19 the the DT cord-346267-l08ld2cy 233 20 development development NN cord-346267-l08ld2cy 233 21 of of IN cord-346267-l08ld2cy 233 22 agriculture agriculture NN cord-346267-l08ld2cy 233 23 allowed allow VBD cord-346267-l08ld2cy 233 24 human human JJ cord-346267-l08ld2cy 233 25 populations population NNS cord-346267-l08ld2cy 233 26 to to TO cord-346267-l08ld2cy 233 27 reach reach VB cord-346267-l08ld2cy 233 28 sizes size NNS cord-346267-l08ld2cy 233 29 necessary necessary JJ cord-346267-l08ld2cy 233 30 to to TO cord-346267-l08ld2cy 233 31 sustain sustain VB cord-346267-l08ld2cy 233 32 an an DT cord-346267-l08ld2cy 233 33 epidemic epidemic NN cord-346267-l08ld2cy 233 34 . . . cord-346267-l08ld2cy 234 1 However however RB cord-346267-l08ld2cy 234 2 , , , cord-346267-l08ld2cy 234 3 the the DT cord-346267-l08ld2cy 234 4 historical historical JJ cord-346267-l08ld2cy 234 5 record record NN cord-346267-l08ld2cy 234 6 of of IN cord-346267-l08ld2cy 234 7 measles measle NNS cord-346267-l08ld2cy 234 8 is be VBZ cord-346267-l08ld2cy 234 9 ambiguous ambiguous JJ cord-346267-l08ld2cy 234 10 until until IN cord-346267-l08ld2cy 234 11 the the DT cord-346267-l08ld2cy 234 12 ninth ninth JJ cord-346267-l08ld2cy 234 13 century century NN cord-346267-l08ld2cy 234 14 , , , cord-346267-l08ld2cy 234 15 when when WRB cord-346267-l08ld2cy 234 16 Rhazes Rhazes NNP cord-346267-l08ld2cy 234 17 ( ( -LRB- cord-346267-l08ld2cy 234 18 a a DT cord-346267-l08ld2cy 234 19 Persian persian JJ cord-346267-l08ld2cy 234 20 physician physician NN cord-346267-l08ld2cy 234 21 ) ) -RRB- cord-346267-l08ld2cy 234 22 outlined outline VBD cord-346267-l08ld2cy 234 23 the the DT cord-346267-l08ld2cy 234 24 criteria criterion NNS cord-346267-l08ld2cy 234 25 for for IN cord-346267-l08ld2cy 234 26 differentiating differentiate VBG cord-346267-l08ld2cy 234 27 between between IN cord-346267-l08ld2cy 234 28 measles measle NNS cord-346267-l08ld2cy 234 29 and and CC cord-346267-l08ld2cy 234 30 smallpox smallpox NN cord-346267-l08ld2cy 234 31 ( ( -LRB- cord-346267-l08ld2cy 234 32 Rāzī Rāzī NNP cord-346267-l08ld2cy 234 33 1848 1848 CD cord-346267-l08ld2cy 234 34 ) ) -RRB- cord-346267-l08ld2cy 234 35 . . . cord-346267-l08ld2cy 235 1 Rhazes Rhazes NNP cord-346267-l08ld2cy 235 2 discusses discuss VBZ cord-346267-l08ld2cy 235 3 both both DT cord-346267-l08ld2cy 235 4 ailments ailment NNS cord-346267-l08ld2cy 235 5 as as IN cord-346267-l08ld2cy 235 6 immemorial immemorial JJ cord-346267-l08ld2cy 235 7 , , , cord-346267-l08ld2cy 235 8 indicating indicate VBG cord-346267-l08ld2cy 235 9 that that IN cord-346267-l08ld2cy 235 10 both both DT cord-346267-l08ld2cy 235 11 diseases disease NNS cord-346267-l08ld2cy 235 12 predated predate VBD cord-346267-l08ld2cy 235 13 him -PRON- PRP cord-346267-l08ld2cy 235 14 by by IN cord-346267-l08ld2cy 235 15 many many JJ cord-346267-l08ld2cy 235 16 generations generation NNS cord-346267-l08ld2cy 235 17 . . . cord-346267-l08ld2cy 236 1 Although although IN cord-346267-l08ld2cy 236 2 there there EX cord-346267-l08ld2cy 236 3 are be VBP cord-346267-l08ld2cy 236 4 historical historical JJ cord-346267-l08ld2cy 236 5 records record NNS cord-346267-l08ld2cy 236 6 of of IN cord-346267-l08ld2cy 236 7 earlier early JJR cord-346267-l08ld2cy 236 8 plagues plague NNS cord-346267-l08ld2cy 236 9 that that WDT cord-346267-l08ld2cy 236 10 could could MD cord-346267-l08ld2cy 236 11 be be VB cord-346267-l08ld2cy 236 12 interpreted interpret VBN cord-346267-l08ld2cy 236 13 as as IN cord-346267-l08ld2cy 236 14 measles measle NNS cord-346267-l08ld2cy 236 15 , , , cord-346267-l08ld2cy 236 16 notably notably RB cord-346267-l08ld2cy 236 17 in in IN cord-346267-l08ld2cy 236 18 eighth eighth JJ cord-346267-l08ld2cy 236 19 century century NN cord-346267-l08ld2cy 236 20 France France NNP cord-346267-l08ld2cy 236 21 during during IN cord-346267-l08ld2cy 236 22 the the DT cord-346267-l08ld2cy 236 23 battle battle NN cord-346267-l08ld2cy 236 24 of of IN cord-346267-l08ld2cy 236 25 Tours Tours NNP cord-346267-l08ld2cy 236 26 ( ( -LRB- cord-346267-l08ld2cy 236 27 Rolleston Rolleston NNP cord-346267-l08ld2cy 236 28 1937 1937 CD cord-346267-l08ld2cy 236 29 ) ) -RRB- cord-346267-l08ld2cy 236 30 , , , cord-346267-l08ld2cy 236 31 it -PRON- PRP cord-346267-l08ld2cy 236 32 is be VBZ cord-346267-l08ld2cy 236 33 also also RB cord-346267-l08ld2cy 236 34 possible possible JJ cord-346267-l08ld2cy 236 35 that that IN cord-346267-l08ld2cy 236 36 this this DT cord-346267-l08ld2cy 236 37 plague plague NN cord-346267-l08ld2cy 236 38 could could MD cord-346267-l08ld2cy 236 39 have have VB cord-346267-l08ld2cy 236 40 been be VBN cord-346267-l08ld2cy 236 41 smallpox smallpox NN cord-346267-l08ld2cy 236 42 or or CC cord-346267-l08ld2cy 236 43 another another DT cord-346267-l08ld2cy 236 44 disease disease NN cord-346267-l08ld2cy 236 45 with with IN cord-346267-l08ld2cy 236 46 similar similar JJ cord-346267-l08ld2cy 236 47 presentation presentation NN cord-346267-l08ld2cy 236 48 . . . cord-346267-l08ld2cy 237 1 Even even RB cord-346267-l08ld2cy 237 2 after after IN cord-346267-l08ld2cy 237 3 Rhazes Rhazes NNPS cord-346267-l08ld2cy 237 4 description description NN cord-346267-l08ld2cy 237 5 , , , cord-346267-l08ld2cy 237 6 Western western JJ cord-346267-l08ld2cy 237 7 medicine medicine NN cord-346267-l08ld2cy 237 8 still still RB cord-346267-l08ld2cy 237 9 confounded confound VBD cord-346267-l08ld2cy 237 10 measles measle NNS cord-346267-l08ld2cy 237 11 , , , cord-346267-l08ld2cy 237 12 smallpox smallpox NN cord-346267-l08ld2cy 237 13 , , , cord-346267-l08ld2cy 237 14 and and CC cord-346267-l08ld2cy 237 15 scarlet scarlet JJ cord-346267-l08ld2cy 237 16 fever fever NN cord-346267-l08ld2cy 237 17 until until IN cord-346267-l08ld2cy 237 18 the the DT cord-346267-l08ld2cy 237 19 17th 17th JJ cord-346267-l08ld2cy 237 20 century century NN cord-346267-l08ld2cy 237 21 ( ( -LRB- cord-346267-l08ld2cy 237 22 Rolleston Rolleston NNP cord-346267-l08ld2cy 237 23 1937 1937 CD cord-346267-l08ld2cy 237 24 ) ) -RRB- cord-346267-l08ld2cy 237 25 . . . cord-346267-l08ld2cy 238 1 In in IN cord-346267-l08ld2cy 238 2 light light NN cord-346267-l08ld2cy 238 3 of of IN cord-346267-l08ld2cy 238 4 this this DT cord-346267-l08ld2cy 238 5 confusion confusion NN cord-346267-l08ld2cy 238 6 , , , cord-346267-l08ld2cy 238 7 the the DT cord-346267-l08ld2cy 238 8 lack lack NN cord-346267-l08ld2cy 238 9 of of IN cord-346267-l08ld2cy 238 10 a a DT cord-346267-l08ld2cy 238 11 definitive definitive JJ cord-346267-l08ld2cy 238 12 description description NN cord-346267-l08ld2cy 238 13 of of IN cord-346267-l08ld2cy 238 14 measles measle NNS cord-346267-l08ld2cy 238 15 before before IN cord-346267-l08ld2cy 238 16 Rhazes Rhazes NNPS cord-346267-l08ld2cy 238 17 ( ( -LRB- cord-346267-l08ld2cy 238 18 e.g. e.g. RB cord-346267-l08ld2cy 238 19 , , , cord-346267-l08ld2cy 238 20 Galen Galen NNP cord-346267-l08ld2cy 238 21 , , , cord-346267-l08ld2cy 238 22 an an DT cord-346267-l08ld2cy 238 23 ancient ancient JJ cord-346267-l08ld2cy 238 24 Roman roman JJ cord-346267-l08ld2cy 238 25 physician physician NN cord-346267-l08ld2cy 238 26 , , , cord-346267-l08ld2cy 238 27 described describe VBD cord-346267-l08ld2cy 238 28 smallpox smallpox NN cord-346267-l08ld2cy 238 29 but but CC cord-346267-l08ld2cy 238 30 not not RB cord-346267-l08ld2cy 238 31 measles measles NN cord-346267-l08ld2cy 238 32 ) ) -RRB- cord-346267-l08ld2cy 238 33 does do VBZ cord-346267-l08ld2cy 238 34 not not RB cord-346267-l08ld2cy 238 35 establish establish VB cord-346267-l08ld2cy 238 36 that that IN cord-346267-l08ld2cy 238 37 the the DT cord-346267-l08ld2cy 238 38 virus virus NN cord-346267-l08ld2cy 238 39 was be VBD cord-346267-l08ld2cy 238 40 absent absent JJ cord-346267-l08ld2cy 238 41 ( ( -LRB- cord-346267-l08ld2cy 238 42 McNeill McNeill NNP cord-346267-l08ld2cy 238 43 1976 1976 CD cord-346267-l08ld2cy 238 44 ) ) -RRB- cord-346267-l08ld2cy 238 45 . . . cord-346267-l08ld2cy 239 1 It -PRON- PRP cord-346267-l08ld2cy 239 2 is be VBZ cord-346267-l08ld2cy 239 3 plausible plausible JJ cord-346267-l08ld2cy 239 4 that that IN cord-346267-l08ld2cy 239 5 MeV MeV NNP cord-346267-l08ld2cy 239 6 might may MD cord-346267-l08ld2cy 239 7 have have VB cord-346267-l08ld2cy 239 8 not not RB cord-346267-l08ld2cy 239 9 entered enter VBN cord-346267-l08ld2cy 239 10 the the DT cord-346267-l08ld2cy 239 11 human human JJ cord-346267-l08ld2cy 239 12 population population NN cord-346267-l08ld2cy 239 13 until until IN cord-346267-l08ld2cy 239 14 the the DT cord-346267-l08ld2cy 239 15 first first JJ cord-346267-l08ld2cy 239 16 millennium millennium NN cord-346267-l08ld2cy 239 17 of of IN cord-346267-l08ld2cy 239 18 the the DT cord-346267-l08ld2cy 239 19 Common Common NNP cord-346267-l08ld2cy 239 20 Era Era NNP cord-346267-l08ld2cy 239 21 , , , cord-346267-l08ld2cy 239 22 as as IN cord-346267-l08ld2cy 239 23 our -PRON- PRP$ cord-346267-l08ld2cy 239 24 analysis analysis NN cord-346267-l08ld2cy 239 25 suggests suggest VBZ cord-346267-l08ld2cy 239 26 . . . cord-346267-l08ld2cy 240 1 Alternatively alternatively RB cord-346267-l08ld2cy 240 2 , , , cord-346267-l08ld2cy 240 3 MeV MeV NNP cord-346267-l08ld2cy 240 4 could could MD cord-346267-l08ld2cy 240 5 have have VB cord-346267-l08ld2cy 240 6 emerged emerge VBN cord-346267-l08ld2cy 240 7 thousands thousand NNS cord-346267-l08ld2cy 240 8 of of IN cord-346267-l08ld2cy 240 9 years year NNS cord-346267-l08ld2cy 240 10 ago ago RB cord-346267-l08ld2cy 240 11 , , , cord-346267-l08ld2cy 240 12 and and CC cord-346267-l08ld2cy 240 13 the the DT cord-346267-l08ld2cy 240 14 evolutionary evolutionary JJ cord-346267-l08ld2cy 240 15 models model NNS cord-346267-l08ld2cy 240 16 employed employ VBN cord-346267-l08ld2cy 240 17 here here RB cord-346267-l08ld2cy 240 18 are be VBP cord-346267-l08ld2cy 240 19 too too RB cord-346267-l08ld2cy 240 20 crude crude JJ cord-346267-l08ld2cy 240 21 to to TO cord-346267-l08ld2cy 240 22 recover recover VB cord-346267-l08ld2cy 240 23 the the DT cord-346267-l08ld2cy 240 24 true true JJ cord-346267-l08ld2cy 240 25 age age NN cord-346267-l08ld2cy 240 26 of of IN cord-346267-l08ld2cy 240 27 the the DT cord-346267-l08ld2cy 240 28 virus virus NN cord-346267-l08ld2cy 240 29 . . . cord-346267-l08ld2cy 241 1 Regardless regardless RB cord-346267-l08ld2cy 241 2 of of IN cord-346267-l08ld2cy 241 3 which which WDT cord-346267-l08ld2cy 241 4 scenario scenario NN cord-346267-l08ld2cy 241 5 is be VBZ cord-346267-l08ld2cy 241 6 correct correct JJ cord-346267-l08ld2cy 241 7 , , , cord-346267-l08ld2cy 241 8 accounting account VBG cord-346267-l08ld2cy 241 9 for for IN cord-346267-l08ld2cy 241 10 variable variable JJ cord-346267-l08ld2cy 241 11 selective selective JJ cord-346267-l08ld2cy 241 12 pressures pressure NNS cord-346267-l08ld2cy 241 13 is be VBZ cord-346267-l08ld2cy 241 14 an an DT cord-346267-l08ld2cy 241 15 important important JJ cord-346267-l08ld2cy 241 16 step step NN cord-346267-l08ld2cy 241 17 in in IN cord-346267-l08ld2cy 241 18 revealing reveal VBG cord-346267-l08ld2cy 241 19 the the DT cord-346267-l08ld2cy 241 20 ancient ancient JJ cord-346267-l08ld2cy 241 21 history history NN cord-346267-l08ld2cy 241 22 of of IN cord-346267-l08ld2cy 241 23 RNA RNA NNP cord-346267-l08ld2cy 241 24 viruses virus NNS cord-346267-l08ld2cy 241 25 . . . cord-346267-l08ld2cy 242 1 The the DT cord-346267-l08ld2cy 242 2 different different JJ cord-346267-l08ld2cy 242 3 timescales timescale NNS cord-346267-l08ld2cy 242 4 affecting affect VBG cord-346267-l08ld2cy 242 5 mutation mutation NN cord-346267-l08ld2cy 242 6 rates rate NNS cord-346267-l08ld2cy 242 7 and and CC cord-346267-l08ld2cy 242 8 substitution substitution NN cord-346267-l08ld2cy 242 9 rates rate NNS cord-346267-l08ld2cy 242 10 make make VBP cord-346267-l08ld2cy 242 11 it -PRON- PRP cord-346267-l08ld2cy 242 12 difficult difficult JJ cord-346267-l08ld2cy 242 13 to to TO cord-346267-l08ld2cy 242 14 extrapolate extrapolate VB cord-346267-l08ld2cy 242 15 population population NN cord-346267-l08ld2cy 242 16 - - HYPH cord-346267-l08ld2cy 242 17 based base VBN cord-346267-l08ld2cy 242 18 estimates estimate NNS cord-346267-l08ld2cy 242 19 of of IN cord-346267-l08ld2cy 242 20 rates rate NNS cord-346267-l08ld2cy 242 21 over over IN cord-346267-l08ld2cy 242 22 a a DT cord-346267-l08ld2cy 242 23 long long JJ cord-346267-l08ld2cy 242 24 evolutionary evolutionary JJ cord-346267-l08ld2cy 242 25 time time NN cord-346267-l08ld2cy 242 26 ( ( -LRB- cord-346267-l08ld2cy 242 27 Ho Ho NNP cord-346267-l08ld2cy 242 28 et et FW cord-346267-l08ld2cy 242 29 al al NNP cord-346267-l08ld2cy 242 30 . . NNP cord-346267-l08ld2cy 242 31 2005 2005 CD cord-346267-l08ld2cy 242 32 ) ) -RRB- cord-346267-l08ld2cy 242 33 ; ; : cord-346267-l08ld2cy 242 34 however however RB cord-346267-l08ld2cy 242 35 , , , cord-346267-l08ld2cy 242 36 it -PRON- PRP cord-346267-l08ld2cy 242 37 is be VBZ cord-346267-l08ld2cy 242 38 unlikely unlikely JJ cord-346267-l08ld2cy 242 39 that that IN cord-346267-l08ld2cy 242 40 purifying purify VBG cord-346267-l08ld2cy 242 41 selection selection NN cord-346267-l08ld2cy 242 42 alone alone RB cord-346267-l08ld2cy 242 43 can can MD cord-346267-l08ld2cy 242 44 account account VB cord-346267-l08ld2cy 242 45 for for IN cord-346267-l08ld2cy 242 46 this this DT cord-346267-l08ld2cy 242 47 difference difference NN cord-346267-l08ld2cy 242 48 between between IN cord-346267-l08ld2cy 242 49 short short JJ cord-346267-l08ld2cy 242 50 - - HYPH cord-346267-l08ld2cy 242 51 and and CC cord-346267-l08ld2cy 242 52 long long JJ cord-346267-l08ld2cy 242 53 - - HYPH cord-346267-l08ld2cy 242 54 term term NN cord-346267-l08ld2cy 242 55 evolutionary evolutionary JJ cord-346267-l08ld2cy 242 56 rates rate NNS cord-346267-l08ld2cy 242 57 ( ( -LRB- cord-346267-l08ld2cy 242 58 Woodhams Woodhams NNP cord-346267-l08ld2cy 242 59 2006 2006 CD cord-346267-l08ld2cy 242 60 ) ) -RRB- cord-346267-l08ld2cy 242 61 . . . cord-346267-l08ld2cy 243 1 Furthermore furthermore RB cord-346267-l08ld2cy 243 2 , , , cord-346267-l08ld2cy 243 3 short short JJ cord-346267-l08ld2cy 243 4 - - HYPH cord-346267-l08ld2cy 243 5 term term NN cord-346267-l08ld2cy 243 6 estimates estimate NNS cord-346267-l08ld2cy 243 7 of of IN cord-346267-l08ld2cy 243 8 viral viral JJ cord-346267-l08ld2cy 243 9 substitution substitution NN cord-346267-l08ld2cy 243 10 rates rate NNS cord-346267-l08ld2cy 243 11 ( ( -LRB- cord-346267-l08ld2cy 243 12 inferred infer VBN cord-346267-l08ld2cy 243 13 from from IN cord-346267-l08ld2cy 243 14 population population NN cord-346267-l08ld2cy 243 15 - - HYPH cord-346267-l08ld2cy 243 16 based base VBN cord-346267-l08ld2cy 243 17 rate rate NN cord-346267-l08ld2cy 243 18 estimates estimate NNS cord-346267-l08ld2cy 243 19 at at IN cord-346267-l08ld2cy 243 20 the the DT cord-346267-l08ld2cy 243 21 tips tip NNS cord-346267-l08ld2cy 243 22 of of IN cord-346267-l08ld2cy 243 23 phylogenies phylogeny NNS cord-346267-l08ld2cy 243 24 ) ) -RRB- cord-346267-l08ld2cy 243 25 are be VBP cord-346267-l08ld2cy 243 26 often often RB cord-346267-l08ld2cy 243 27 found find VBN cord-346267-l08ld2cy 243 28 to to TO cord-346267-l08ld2cy 243 29 be be VB cord-346267-l08ld2cy 243 30 several several JJ cord-346267-l08ld2cy 243 31 orders order NNS cord-346267-l08ld2cy 243 32 of of IN cord-346267-l08ld2cy 243 33 magnitude magnitude NN cord-346267-l08ld2cy 243 34 faster fast RBR cord-346267-l08ld2cy 243 35 than than IN cord-346267-l08ld2cy 243 36 long long JJ cord-346267-l08ld2cy 243 37 - - HYPH cord-346267-l08ld2cy 243 38 term term NN cord-346267-l08ld2cy 243 39 estimates estimate NNS cord-346267-l08ld2cy 243 40 of of IN cord-346267-l08ld2cy 243 41 substitution substitution NN cord-346267-l08ld2cy 243 42 rates rate NNS cord-346267-l08ld2cy 243 43 ( ( -LRB- cord-346267-l08ld2cy 243 44 inferred infer VBN cord-346267-l08ld2cy 243 45 from from IN cord-346267-l08ld2cy 243 46 external external JJ cord-346267-l08ld2cy 243 47 calibrations calibration NNS cord-346267-l08ld2cy 243 48 located locate VBN cord-346267-l08ld2cy 243 49 deeper deeply RBR cord-346267-l08ld2cy 243 50 in in IN cord-346267-l08ld2cy 243 51 the the DT cord-346267-l08ld2cy 243 52 phylogeny phylogeny NN cord-346267-l08ld2cy 243 53 ) ) -RRB- cord-346267-l08ld2cy 243 54 . . . cord-346267-l08ld2cy 244 1 This this DT cord-346267-l08ld2cy 244 2 inconsistency inconsistency NN cord-346267-l08ld2cy 244 3 has have VBZ cord-346267-l08ld2cy 244 4 been be VBN cord-346267-l08ld2cy 244 5 reported report VBN cord-346267-l08ld2cy 244 6 for for IN cord-346267-l08ld2cy 244 7 hepatitis hepatitis NN cord-346267-l08ld2cy 244 8 B B NNP cord-346267-l08ld2cy 244 9 virus virus NN cord-346267-l08ld2cy 244 10 ( ( -LRB- cord-346267-l08ld2cy 244 11 Zhou Zhou NNP cord-346267-l08ld2cy 244 12 and and CC cord-346267-l08ld2cy 244 13 Holmes Holmes NNP cord-346267-l08ld2cy 244 14 2007 2007 CD cord-346267-l08ld2cy 244 15 ; ; : cord-346267-l08ld2cy 244 16 Gilbert Gilbert NNP cord-346267-l08ld2cy 244 17 and and CC cord-346267-l08ld2cy 244 18 Feschotte Feschotte NNP cord-346267-l08ld2cy 244 19 2010 2010 CD cord-346267-l08ld2cy 244 20 ) ) -RRB- cord-346267-l08ld2cy 244 21 and and CC cord-346267-l08ld2cy 244 22 simian simian JJ cord-346267-l08ld2cy 244 23 immunodeficiency immunodeficiency NN cord-346267-l08ld2cy 244 24 virus virus NN cord-346267-l08ld2cy 244 25 ( ( -LRB- cord-346267-l08ld2cy 244 26 Wertheim Wertheim NNP cord-346267-l08ld2cy 244 27 and and CC cord-346267-l08ld2cy 244 28 Worobey Worobey NNP cord-346267-l08ld2cy 244 29 2009a 2009a CD cord-346267-l08ld2cy 244 30 ; ; : cord-346267-l08ld2cy 244 31 Worobey Worobey NNP cord-346267-l08ld2cy 244 32 et et NNP cord-346267-l08ld2cy 244 33 al al NNP cord-346267-l08ld2cy 244 34 . . NNP cord-346267-l08ld2cy 244 35 2010 2010 CD cord-346267-l08ld2cy 244 36 ) ) -RRB- cord-346267-l08ld2cy 244 37 , , . cord-346267-l08ld2cy 245 1 suggesting suggest VBG cord-346267-l08ld2cy 245 2 that that IN cord-346267-l08ld2cy 245 3 long long JJ cord-346267-l08ld2cy 245 4 - - HYPH cord-346267-l08ld2cy 245 5 term term NN cord-346267-l08ld2cy 245 6 substitution substitution NN cord-346267-l08ld2cy 245 7 rates rate NNS cord-346267-l08ld2cy 245 8 can can MD cord-346267-l08ld2cy 245 9 be be VB cord-346267-l08ld2cy 245 10 orders order NNS cord-346267-l08ld2cy 245 11 of of IN cord-346267-l08ld2cy 245 12 magnitude magnitude NN cord-346267-l08ld2cy 245 13 lower low JJR cord-346267-l08ld2cy 245 14 than than IN cord-346267-l08ld2cy 245 15 short short JJ cord-346267-l08ld2cy 245 16 - - HYPH cord-346267-l08ld2cy 245 17 term term NN cord-346267-l08ld2cy 245 18 substitution substitution NN cord-346267-l08ld2cy 245 19 rates rate NNS cord-346267-l08ld2cy 245 20 . . . cord-346267-l08ld2cy 246 1 An an DT cord-346267-l08ld2cy 246 2 alternative alternative NN cord-346267-l08ld2cy 246 3 , , , cord-346267-l08ld2cy 246 4 and and CC cord-346267-l08ld2cy 246 5 more more RBR cord-346267-l08ld2cy 246 6 parsimonious parsimonious JJ cord-346267-l08ld2cy 246 7 , , , cord-346267-l08ld2cy 246 8 explanation explanation NN cord-346267-l08ld2cy 246 9 is be VBZ cord-346267-l08ld2cy 246 10 that that IN cord-346267-l08ld2cy 246 11 a a DT cord-346267-l08ld2cy 246 12 single single JJ cord-346267-l08ld2cy 246 13 substitution substitution NN cord-346267-l08ld2cy 246 14 rate rate NN cord-346267-l08ld2cy 246 15 ( ( -LRB- cord-346267-l08ld2cy 246 16 albeit albeit IN cord-346267-l08ld2cy 246 17 one one CD cord-346267-l08ld2cy 246 18 possibly possibly RB cord-346267-l08ld2cy 246 19 slower slow JJR cord-346267-l08ld2cy 246 20 than than IN cord-346267-l08ld2cy 246 21 that that DT cord-346267-l08ld2cy 246 22 inferred infer VBN cord-346267-l08ld2cy 246 23 using use VBG cord-346267-l08ld2cy 246 24 short short JJ cord-346267-l08ld2cy 246 25 - - HYPH cord-346267-l08ld2cy 246 26 term term NN cord-346267-l08ld2cy 246 27 population population NN cord-346267-l08ld2cy 246 28 - - HYPH cord-346267-l08ld2cy 246 29 based base VBN cord-346267-l08ld2cy 246 30 data datum NNS cord-346267-l08ld2cy 246 31 ) ) -RRB- cord-346267-l08ld2cy 246 32 predominates predominate VBZ cord-346267-l08ld2cy 246 33 throughout throughout IN cord-346267-l08ld2cy 246 34 the the DT cord-346267-l08ld2cy 246 35 history history NN cord-346267-l08ld2cy 246 36 of of IN cord-346267-l08ld2cy 246 37 these these DT cord-346267-l08ld2cy 246 38 viruses virus NNS cord-346267-l08ld2cy 246 39 . . . cord-346267-l08ld2cy 247 1 And and CC cord-346267-l08ld2cy 247 2 our -PRON- PRP$ cord-346267-l08ld2cy 247 3 inability inability NN cord-346267-l08ld2cy 247 4 to to TO cord-346267-l08ld2cy 247 5 accurately accurately RB cord-346267-l08ld2cy 247 6 estimate estimate VB cord-346267-l08ld2cy 247 7 branch branch NN cord-346267-l08ld2cy 247 8 lengths length NNS cord-346267-l08ld2cy 247 9 creates create VBZ cord-346267-l08ld2cy 247 10 the the DT cord-346267-l08ld2cy 247 11 appearance appearance NN cord-346267-l08ld2cy 247 12 of of IN cord-346267-l08ld2cy 247 13 dramatically dramatically RB cord-346267-l08ld2cy 247 14 lower low JJR cord-346267-l08ld2cy 247 15 substitution substitution NN cord-346267-l08ld2cy 247 16 rates rate NNS cord-346267-l08ld2cy 247 17 deep deep JJ cord-346267-l08ld2cy 247 18 in in IN cord-346267-l08ld2cy 247 19 the the DT cord-346267-l08ld2cy 247 20 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 247 21 tree tree NN cord-346267-l08ld2cy 247 22 . . . cord-346267-l08ld2cy 248 1 Although although IN cord-346267-l08ld2cy 248 2 the the DT cord-346267-l08ld2cy 248 3 methods method NNS cord-346267-l08ld2cy 248 4 presented present VBN cord-346267-l08ld2cy 248 5 here here RB cord-346267-l08ld2cy 248 6 do do VBP cord-346267-l08ld2cy 248 7 not not RB cord-346267-l08ld2cy 248 8 correct correct VB cord-346267-l08ld2cy 248 9 for for IN cord-346267-l08ld2cy 248 10 branch branch NN cord-346267-l08ld2cy 248 11 lengths length NNS cord-346267-l08ld2cy 248 12 on on IN cord-346267-l08ld2cy 248 13 the the DT cord-346267-l08ld2cy 248 14 order order NN cord-346267-l08ld2cy 248 15 needed need VBD cord-346267-l08ld2cy 248 16 to to TO cord-346267-l08ld2cy 248 17 reconcile reconcile VB cord-346267-l08ld2cy 248 18 short short JJ cord-346267-l08ld2cy 248 19 - - HYPH cord-346267-l08ld2cy 248 20 term term NN cord-346267-l08ld2cy 248 21 substitution substitution NN cord-346267-l08ld2cy 248 22 rates rate NNS cord-346267-l08ld2cy 248 23 with with IN cord-346267-l08ld2cy 248 24 deep deep JJ cord-346267-l08ld2cy 248 25 calibrations calibration NNS cord-346267-l08ld2cy 248 26 , , , cord-346267-l08ld2cy 248 27 they -PRON- PRP cord-346267-l08ld2cy 248 28 provide provide VBP cord-346267-l08ld2cy 248 29 a a DT cord-346267-l08ld2cy 248 30 glimpse glimpse NN cord-346267-l08ld2cy 248 31 at at IN cord-346267-l08ld2cy 248 32 the the DT cord-346267-l08ld2cy 248 33 missing missing JJ cord-346267-l08ld2cy 248 34 evolution evolution NN cord-346267-l08ld2cy 248 35 that that IN cord-346267-l08ld2cy 248 36 most most JJS cord-346267-l08ld2cy 248 37 current current JJ cord-346267-l08ld2cy 248 38 methods method NNS cord-346267-l08ld2cy 248 39 of of IN cord-346267-l08ld2cy 248 40 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 248 41 inference inference NN cord-346267-l08ld2cy 248 42 fail fail VBP cord-346267-l08ld2cy 248 43 to to TO cord-346267-l08ld2cy 248 44 capture capture VB cord-346267-l08ld2cy 248 45 . . . cord-346267-l08ld2cy 249 1 Although although IN cord-346267-l08ld2cy 249 2 we -PRON- PRP cord-346267-l08ld2cy 249 3 are be VBP cord-346267-l08ld2cy 249 4 unable unable JJ cord-346267-l08ld2cy 249 5 to to TO cord-346267-l08ld2cy 249 6 provide provide VB cord-346267-l08ld2cy 249 7 a a DT cord-346267-l08ld2cy 249 8 full full JJ cord-346267-l08ld2cy 249 9 remedy remedy NN cord-346267-l08ld2cy 249 10 to to IN cord-346267-l08ld2cy 249 11 the the DT cord-346267-l08ld2cy 249 12 problem problem NN cord-346267-l08ld2cy 249 13 of of IN cord-346267-l08ld2cy 249 14 underestimated underestimated JJ cord-346267-l08ld2cy 249 15 branch branch NN cord-346267-l08ld2cy 249 16 lengths length NNS cord-346267-l08ld2cy 249 17 due due JJ cord-346267-l08ld2cy 249 18 to to IN cord-346267-l08ld2cy 249 19 purifying purify VBG cord-346267-l08ld2cy 249 20 selection selection NN cord-346267-l08ld2cy 249 21 , , , cord-346267-l08ld2cy 249 22 our -PRON- PRP$ cord-346267-l08ld2cy 249 23 results result NNS cord-346267-l08ld2cy 249 24 do do VBP cord-346267-l08ld2cy 249 25 point point NN cord-346267-l08ld2cy 249 26 to to IN cord-346267-l08ld2cy 249 27 a a DT cord-346267-l08ld2cy 249 28 couple couple NN cord-346267-l08ld2cy 249 29 of of IN cord-346267-l08ld2cy 249 30 guidelines guideline NNS cord-346267-l08ld2cy 249 31 when when WRB cord-346267-l08ld2cy 249 32 inferring infer VBG cord-346267-l08ld2cy 249 33 tMRCAs tmrcas CD cord-346267-l08ld2cy 249 34 in in IN cord-346267-l08ld2cy 249 35 RNA RNA NNP cord-346267-l08ld2cy 249 36 viruses virus NNS cord-346267-l08ld2cy 249 37 . . . cord-346267-l08ld2cy 250 1 First first RB cord-346267-l08ld2cy 250 2 , , , cord-346267-l08ld2cy 250 3 the the DT cord-346267-l08ld2cy 250 4 inference inference NN cord-346267-l08ld2cy 250 5 of of IN cord-346267-l08ld2cy 250 6 substantially substantially RB cord-346267-l08ld2cy 250 7 different different JJ cord-346267-l08ld2cy 250 8 selective selective JJ cord-346267-l08ld2cy 250 9 regimes regime NNS cord-346267-l08ld2cy 250 10 on on IN cord-346267-l08ld2cy 250 11 longer long JJR cord-346267-l08ld2cy 250 12 internal internal JJ cord-346267-l08ld2cy 250 13 branches branch NNS cord-346267-l08ld2cy 250 14 compared compare VBN cord-346267-l08ld2cy 250 15 with with IN cord-346267-l08ld2cy 250 16 shorter short JJR cord-346267-l08ld2cy 250 17 shallower shallow JJR cord-346267-l08ld2cy 250 18 branches branch NNS cord-346267-l08ld2cy 250 19 ( ( -LRB- cord-346267-l08ld2cy 250 20 e.g. e.g. RB cord-346267-l08ld2cy 250 21 , , , cord-346267-l08ld2cy 250 22 using use VBG cord-346267-l08ld2cy 250 23 Lineage+Dual lineage+dual JJ cord-346267-l08ld2cy 250 24 , , , cord-346267-l08ld2cy 250 25 , , , cord-346267-l08ld2cy 250 26 or or CC cord-346267-l08ld2cy 250 27 Free Free NNP cord-346267-l08ld2cy 250 28 Ratio Ratio NNP cord-346267-l08ld2cy 250 29 , , , cord-346267-l08ld2cy 250 30 Yang Yang NNP cord-346267-l08ld2cy 250 31 1998 1998 CD cord-346267-l08ld2cy 250 32 , , , cord-346267-l08ld2cy 250 33 codon codon NN cord-346267-l08ld2cy 250 34 models model NNS cord-346267-l08ld2cy 250 35 ) ) -RRB- cord-346267-l08ld2cy 250 36 appears appear VBZ cord-346267-l08ld2cy 250 37 to to TO cord-346267-l08ld2cy 250 38 be be VB cord-346267-l08ld2cy 250 39 a a DT cord-346267-l08ld2cy 250 40 sign sign NN cord-346267-l08ld2cy 250 41 that that IN cord-346267-l08ld2cy 250 42 older old JJR cord-346267-l08ld2cy 250 43 tMRCAs tmrcas PRP cord-346267-l08ld2cy 250 44 may may MD cord-346267-l08ld2cy 250 45 be be VB cord-346267-l08ld2cy 250 46 underestimated underestimate VBN cord-346267-l08ld2cy 250 47 . . . cord-346267-l08ld2cy 251 1 Second second RB cord-346267-l08ld2cy 251 2 , , , cord-346267-l08ld2cy 251 3 if if IN cord-346267-l08ld2cy 251 4 the the DT cord-346267-l08ld2cy 251 5 synonymous synonymous JJ cord-346267-l08ld2cy 251 6 substitution substitution NN cord-346267-l08ld2cy 251 7 rate rate NN cord-346267-l08ld2cy 251 8 approaches approach VBZ cord-346267-l08ld2cy 251 9 saturation saturation VBP cord-346267-l08ld2cy 251 10 along along IN cord-346267-l08ld2cy 251 11 a a DT cord-346267-l08ld2cy 251 12 branch branch NN cord-346267-l08ld2cy 251 13 or or CC cord-346267-l08ld2cy 251 14 group group NN cord-346267-l08ld2cy 251 15 of of IN cord-346267-l08ld2cy 251 16 branches branch NNS cord-346267-l08ld2cy 251 17 , , , cord-346267-l08ld2cy 251 18 it -PRON- PRP cord-346267-l08ld2cy 251 19 is be VBZ cord-346267-l08ld2cy 251 20 likely likely JJ cord-346267-l08ld2cy 251 21 that that IN cord-346267-l08ld2cy 251 22 the the DT cord-346267-l08ld2cy 251 23 tMRCA tmrca NN cord-346267-l08ld2cy 251 24 can can MD cord-346267-l08ld2cy 251 25 not not RB cord-346267-l08ld2cy 251 26 be be VB cord-346267-l08ld2cy 251 27 reliably reliably RB cord-346267-l08ld2cy 251 28 inferred infer VBN cord-346267-l08ld2cy 251 29 . . . cord-346267-l08ld2cy 252 1 If if IN cord-346267-l08ld2cy 252 2 either either DT cord-346267-l08ld2cy 252 3 of of IN cord-346267-l08ld2cy 252 4 these these DT cord-346267-l08ld2cy 252 5 patterns pattern NNS cord-346267-l08ld2cy 252 6 is be VBZ cord-346267-l08ld2cy 252 7 encountered encounter VBN cord-346267-l08ld2cy 252 8 , , , cord-346267-l08ld2cy 252 9 the the DT cord-346267-l08ld2cy 252 10 inferred infer VBN cord-346267-l08ld2cy 252 11 tMRCA tmrca NN cord-346267-l08ld2cy 252 12 estimates estimate NNS cord-346267-l08ld2cy 252 13 should should MD cord-346267-l08ld2cy 252 14 be be VB cord-346267-l08ld2cy 252 15 interpreted interpret VBN cord-346267-l08ld2cy 252 16 with with IN cord-346267-l08ld2cy 252 17 caution caution NN cord-346267-l08ld2cy 252 18 . . . cord-346267-l08ld2cy 253 1 Rapid rapid JJ cord-346267-l08ld2cy 253 2 advances advance NNS cord-346267-l08ld2cy 253 3 in in IN cord-346267-l08ld2cy 253 4 cheap cheap JJ cord-346267-l08ld2cy 253 5 and and CC cord-346267-l08ld2cy 253 6 accessible accessible JJ cord-346267-l08ld2cy 253 7 computational computational JJ cord-346267-l08ld2cy 253 8 power power NN cord-346267-l08ld2cy 253 9 will will MD cord-346267-l08ld2cy 253 10 undoubtedly undoubtedly RB cord-346267-l08ld2cy 253 11 move move VB cord-346267-l08ld2cy 253 12 the the DT cord-346267-l08ld2cy 253 13 fields field NNS cord-346267-l08ld2cy 253 14 of of IN cord-346267-l08ld2cy 253 15 paleovirology paleovirology NN cord-346267-l08ld2cy 253 16 and and CC cord-346267-l08ld2cy 253 17 molecular molecular JJ cord-346267-l08ld2cy 253 18 dating date VBG cord-346267-l08ld2cy 253 19 towards towards IN cord-346267-l08ld2cy 253 20 increasingly increasingly RB cord-346267-l08ld2cy 253 21 more more RBR cord-346267-l08ld2cy 253 22 realistic realistic JJ cord-346267-l08ld2cy 253 23 evolutionary evolutionary JJ cord-346267-l08ld2cy 253 24 models model NNS cord-346267-l08ld2cy 253 25 that that WDT cord-346267-l08ld2cy 253 26 account account VBP cord-346267-l08ld2cy 253 27 for for IN cord-346267-l08ld2cy 253 28 variation variation NN cord-346267-l08ld2cy 253 29 in in IN cord-346267-l08ld2cy 253 30 selective selective JJ cord-346267-l08ld2cy 253 31 regimes regime NNS cord-346267-l08ld2cy 253 32 ( ( -LRB- cord-346267-l08ld2cy 253 33 Suchard Suchard NNP cord-346267-l08ld2cy 253 34 and and CC cord-346267-l08ld2cy 253 35 Rambaut Rambaut NNP cord-346267-l08ld2cy 253 36 2009 2009 CD cord-346267-l08ld2cy 253 37 ) ) -RRB- cord-346267-l08ld2cy 253 38 . . . cord-346267-l08ld2cy 254 1 Our -PRON- PRP$ cord-346267-l08ld2cy 254 2 results result NNS cord-346267-l08ld2cy 254 3 provide provide VBP cord-346267-l08ld2cy 254 4 compelling compelling JJ cord-346267-l08ld2cy 254 5 evidence evidence NN cord-346267-l08ld2cy 254 6 that that IN cord-346267-l08ld2cy 254 7 such such PDT cord-346267-l08ld2cy 254 8 a a DT cord-346267-l08ld2cy 254 9 movement movement NN cord-346267-l08ld2cy 254 10 should should MD cord-346267-l08ld2cy 254 11 be be VB cord-346267-l08ld2cy 254 12 accelerated accelerate VBN cord-346267-l08ld2cy 254 13 . . . cord-346267-l08ld2cy 255 1 In in IN cord-346267-l08ld2cy 255 2 this this DT cord-346267-l08ld2cy 255 3 study study NN cord-346267-l08ld2cy 255 4 , , , cord-346267-l08ld2cy 255 5 we -PRON- PRP cord-346267-l08ld2cy 255 6 demonstrate demonstrate VBP cord-346267-l08ld2cy 255 7 the the DT cord-346267-l08ld2cy 255 8 need need NN cord-346267-l08ld2cy 255 9 to to TO cord-346267-l08ld2cy 255 10 include include VB cord-346267-l08ld2cy 255 11 relevant relevant JJ cord-346267-l08ld2cy 255 12 biological biological JJ cord-346267-l08ld2cy 255 13 and and CC cord-346267-l08ld2cy 255 14 evolutionary evolutionary JJ cord-346267-l08ld2cy 255 15 forces force NNS cord-346267-l08ld2cy 255 16 in in IN cord-346267-l08ld2cy 255 17 substitution substitution NN cord-346267-l08ld2cy 255 18 models model NNS cord-346267-l08ld2cy 255 19 . . . cord-346267-l08ld2cy 256 1 It -PRON- PRP cord-346267-l08ld2cy 256 2 was be VBD cord-346267-l08ld2cy 256 3 not not RB cord-346267-l08ld2cy 256 4 until until IN cord-346267-l08ld2cy 256 5 we -PRON- PRP cord-346267-l08ld2cy 256 6 permitted permit VBD cord-346267-l08ld2cy 256 7 different different JJ cord-346267-l08ld2cy 256 8 evolutionary evolutionary JJ cord-346267-l08ld2cy 256 9 regimes regime NNS cord-346267-l08ld2cy 256 10 for for IN cord-346267-l08ld2cy 256 11 different different JJ cord-346267-l08ld2cy 256 12 branches branch NNS cord-346267-l08ld2cy 256 13 in in IN cord-346267-l08ld2cy 256 14 the the DT cord-346267-l08ld2cy 256 15 tree tree NN cord-346267-l08ld2cy 256 16 that that WDT cord-346267-l08ld2cy 256 17 we -PRON- PRP cord-346267-l08ld2cy 256 18 saw see VBD cord-346267-l08ld2cy 256 19 the the DT cord-346267-l08ld2cy 256 20 most most RBS cord-346267-l08ld2cy 256 21 dramatic dramatic JJ cord-346267-l08ld2cy 256 22 changes change NNS cord-346267-l08ld2cy 256 23 in in IN cord-346267-l08ld2cy 256 24 branch branch NN cord-346267-l08ld2cy 256 25 length length NN cord-346267-l08ld2cy 256 26 estimates estimate NNS cord-346267-l08ld2cy 256 27 . . . cord-346267-l08ld2cy 257 1 Current current JJ cord-346267-l08ld2cy 257 2 codon codon NN cord-346267-l08ld2cy 257 3 models model NNS cord-346267-l08ld2cy 257 4 take take VBP cord-346267-l08ld2cy 257 5 a a DT cord-346267-l08ld2cy 257 6 very very RB cord-346267-l08ld2cy 257 7 simplistic simplistic JJ cord-346267-l08ld2cy 257 8 mechanistic mechanistic JJ cord-346267-l08ld2cy 257 9 view view NN cord-346267-l08ld2cy 257 10 of of IN cord-346267-l08ld2cy 257 11 the the DT cord-346267-l08ld2cy 257 12 action action NN cord-346267-l08ld2cy 257 13 of of IN cord-346267-l08ld2cy 257 14 purifying purify VBG cord-346267-l08ld2cy 257 15 selection selection NN cord-346267-l08ld2cy 257 16 , , , cord-346267-l08ld2cy 257 17 and and CC cord-346267-l08ld2cy 257 18 we -PRON- PRP cord-346267-l08ld2cy 257 19 expect expect VBP cord-346267-l08ld2cy 257 20 that that IN cord-346267-l08ld2cy 257 21 incorporating incorporate VBG cord-346267-l08ld2cy 257 22 processes process NNS cord-346267-l08ld2cy 257 23 such such JJ cord-346267-l08ld2cy 257 24 as as IN cord-346267-l08ld2cy 257 25 directional directional JJ cord-346267-l08ld2cy 257 26 selection selection NN cord-346267-l08ld2cy 257 27 ( ( -LRB- cord-346267-l08ld2cy 257 28 Seoighe Seoighe NNP cord-346267-l08ld2cy 257 29 et et NNP cord-346267-l08ld2cy 257 30 al al NNP cord-346267-l08ld2cy 257 31 . . NNP cord-346267-l08ld2cy 257 32 2007 2007 CD cord-346267-l08ld2cy 257 33 ) ) -RRB- cord-346267-l08ld2cy 257 34 , , , cord-346267-l08ld2cy 257 35 toggling toggle VBG cord-346267-l08ld2cy 257 36 selection selection NN cord-346267-l08ld2cy 257 37 ( ( -LRB- cord-346267-l08ld2cy 257 38 Delport Delport NNP cord-346267-l08ld2cy 257 39 et et NNP cord-346267-l08ld2cy 257 40 al al NNP cord-346267-l08ld2cy 257 41 . . NNP cord-346267-l08ld2cy 257 42 2008 2008 CD cord-346267-l08ld2cy 257 43 ) ) -RRB- cord-346267-l08ld2cy 257 44 , , , cord-346267-l08ld2cy 257 45 and and CC cord-346267-l08ld2cy 257 46 residue residue NN cord-346267-l08ld2cy 257 47 - - HYPH cord-346267-l08ld2cy 257 48 specific specific JJ cord-346267-l08ld2cy 257 49 ( ( -LRB- cord-346267-l08ld2cy 257 50 Doron Doron NNP cord-346267-l08ld2cy 257 51 - - HYPH cord-346267-l08ld2cy 257 52 Faigenboim Faigenboim NNP cord-346267-l08ld2cy 257 53 and and CC cord-346267-l08ld2cy 257 54 Pupko Pupko NNP cord-346267-l08ld2cy 257 55 2007 2007 CD cord-346267-l08ld2cy 257 56 ) ) -RRB- cord-346267-l08ld2cy 257 57 or or CC cord-346267-l08ld2cy 257 58 site site NN cord-346267-l08ld2cy 257 59 - - HYPH cord-346267-l08ld2cy 257 60 specific specific JJ cord-346267-l08ld2cy 257 61 substitution substitution NN cord-346267-l08ld2cy 257 62 ( ( -LRB- cord-346267-l08ld2cy 257 63 Lartillot Lartillot NNP cord-346267-l08ld2cy 257 64 and and CC cord-346267-l08ld2cy 257 65 Philippe Philippe NNP cord-346267-l08ld2cy 257 66 2004 2004 CD cord-346267-l08ld2cy 257 67 ) ) -RRB- cord-346267-l08ld2cy 257 68 biases bias NNS cord-346267-l08ld2cy 257 69 can can MD cord-346267-l08ld2cy 257 70 further further RB cord-346267-l08ld2cy 257 71 refine refine VB cord-346267-l08ld2cy 257 72 tMRCA tmrca NN cord-346267-l08ld2cy 257 73 estimates estimate NNS cord-346267-l08ld2cy 257 74 . . . cord-346267-l08ld2cy 258 1 It -PRON- PRP cord-346267-l08ld2cy 258 2 is be VBZ cord-346267-l08ld2cy 258 3 still still RB cord-346267-l08ld2cy 258 4 unclear unclear JJ cord-346267-l08ld2cy 258 5 how how WRB cord-346267-l08ld2cy 258 6 correcting correct VBG cord-346267-l08ld2cy 258 7 for for IN cord-346267-l08ld2cy 258 8 the the DT cord-346267-l08ld2cy 258 9 effects effect NNS cord-346267-l08ld2cy 258 10 of of IN cord-346267-l08ld2cy 258 11 purifying purify VBG cord-346267-l08ld2cy 258 12 selection selection NN cord-346267-l08ld2cy 258 13 may may MD cord-346267-l08ld2cy 258 14 affect affect VB cord-346267-l08ld2cy 258 15 dating dating NN cord-346267-l08ld2cy 258 16 and and CC cord-346267-l08ld2cy 258 17 branch branch NN cord-346267-l08ld2cy 258 18 length length NN cord-346267-l08ld2cy 258 19 estimation estimation NN cord-346267-l08ld2cy 258 20 in in IN cord-346267-l08ld2cy 258 21 other other JJ cord-346267-l08ld2cy 258 22 types type NNS cord-346267-l08ld2cy 258 23 of of IN cord-346267-l08ld2cy 258 24 viruses virus NNS cord-346267-l08ld2cy 258 25 ( ( -LRB- cord-346267-l08ld2cy 258 26 e.g. e.g. RB cord-346267-l08ld2cy 258 27 , , , cord-346267-l08ld2cy 258 28 DNA dna NN cord-346267-l08ld2cy 258 29 viruses virus NNS cord-346267-l08ld2cy 258 30 ) ) -RRB- cord-346267-l08ld2cy 258 31 and and CC cord-346267-l08ld2cy 258 32 cellular cellular JJ cord-346267-l08ld2cy 258 33 organisms organism NNS cord-346267-l08ld2cy 258 34 . . . cord-346267-l08ld2cy 259 1 The the DT cord-346267-l08ld2cy 259 2 bias bias NN cord-346267-l08ld2cy 259 3 we -PRON- PRP cord-346267-l08ld2cy 259 4 observed observe VBD cord-346267-l08ld2cy 259 5 in in IN cord-346267-l08ld2cy 259 6 the the DT cord-346267-l08ld2cy 259 7 estimation estimation NN cord-346267-l08ld2cy 259 8 of of IN cord-346267-l08ld2cy 259 9 branch branch NN cord-346267-l08ld2cy 259 10 lengths length NNS cord-346267-l08ld2cy 259 11 appears appear VBZ cord-346267-l08ld2cy 259 12 to to TO cord-346267-l08ld2cy 259 13 be be VB cord-346267-l08ld2cy 259 14 related relate VBN cord-346267-l08ld2cy 259 15 to to IN cord-346267-l08ld2cy 259 16 the the DT cord-346267-l08ld2cy 259 17 intensity intensity NN cord-346267-l08ld2cy 259 18 of of IN cord-346267-l08ld2cy 259 19 selection selection NN cord-346267-l08ld2cy 259 20 and and CC cord-346267-l08ld2cy 259 21 the the DT cord-346267-l08ld2cy 259 22 length length NN cord-346267-l08ld2cy 259 23 of of IN cord-346267-l08ld2cy 259 24 the the DT cord-346267-l08ld2cy 259 25 branches branch NNS cord-346267-l08ld2cy 259 26 in in IN cord-346267-l08ld2cy 259 27 question question NN cord-346267-l08ld2cy 259 28 . . . cord-346267-l08ld2cy 260 1 The the DT cord-346267-l08ld2cy 260 2 genome genome NN cord-346267-l08ld2cy 260 3 sequence sequence NN cord-346267-l08ld2cy 260 4 of of IN cord-346267-l08ld2cy 260 5 the the DT cord-346267-l08ld2cy 260 6 virulent virulent JJ cord-346267-l08ld2cy 260 7 Kabete Kabete NNP cord-346267-l08ld2cy 260 8 ' ' `` cord-346267-l08ld2cy 260 9 O o UH cord-346267-l08ld2cy 260 10 ' ' '' cord-346267-l08ld2cy 260 11 strain strain NN cord-346267-l08ld2cy 260 12 of of IN cord-346267-l08ld2cy 260 13 rinderpest rinderpest NN cord-346267-l08ld2cy 260 14 virus virus NN cord-346267-l08ld2cy 260 15 : : : cord-346267-l08ld2cy 260 16 comparison comparison NN cord-346267-l08ld2cy 260 17 with with IN cord-346267-l08ld2cy 260 18 the the DT cord-346267-l08ld2cy 260 19 derived derived JJ cord-346267-l08ld2cy 260 20 vaccine vaccine NN cord-346267-l08ld2cy 260 21 The the DT cord-346267-l08ld2cy 260 22 origin origin NN cord-346267-l08ld2cy 260 23 of of IN cord-346267-l08ld2cy 260 24 European european JJ cord-346267-l08ld2cy 260 25 cattle cattle NNS cord-346267-l08ld2cy 260 26 : : : cord-346267-l08ld2cy 260 27 evidence evidence NN cord-346267-l08ld2cy 260 28 from from IN cord-346267-l08ld2cy 260 29 modern modern JJ cord-346267-l08ld2cy 260 30 and and CC cord-346267-l08ld2cy 260 31 ancient ancient JJ cord-346267-l08ld2cy 260 32 DNA dna NN cord-346267-l08ld2cy 260 33 Pacing pace VBG cord-346267-l08ld2cy 260 34 a a DT cord-346267-l08ld2cy 260 35 small small JJ cord-346267-l08ld2cy 260 36 cage cage NN cord-346267-l08ld2cy 260 37 : : : cord-346267-l08ld2cy 260 38 mutation mutation NN cord-346267-l08ld2cy 260 39 and and CC cord-346267-l08ld2cy 260 40 RNA RNA NNP cord-346267-l08ld2cy 260 41 viruses virus NNS cord-346267-l08ld2cy 260 42 Unexpected unexpected JJ cord-346267-l08ld2cy 260 43 inheritance inheritance NN cord-346267-l08ld2cy 260 44 : : : cord-346267-l08ld2cy 260 45 multiple multiple JJ cord-346267-l08ld2cy 260 46 integrations integration NNS cord-346267-l08ld2cy 260 47 of of IN cord-346267-l08ld2cy 260 48 ancient ancient JJ cord-346267-l08ld2cy 260 49 bornavirus bornavirus NN cord-346267-l08ld2cy 260 50 and and CC cord-346267-l08ld2cy 260 51 ebolavirus/ ebolavirus/ NN cord-346267-l08ld2cy 260 52 marburgvirus marburgvirus JJ cord-346267-l08ld2cy 260 53 sequences sequence NNS cord-346267-l08ld2cy 260 54 in in IN cord-346267-l08ld2cy 260 55 vertebrate vertebrate JJ cord-346267-l08ld2cy 260 56 genomes genome NNS cord-346267-l08ld2cy 260 57 Measles measles VBP cord-346267-l08ld2cy 260 58 endemicity endemicity NN cord-346267-l08ld2cy 260 59 in in IN cord-346267-l08ld2cy 260 60 insular insular JJ cord-346267-l08ld2cy 260 61 populations population NNS cord-346267-l08ld2cy 260 62 : : : cord-346267-l08ld2cy 260 63 critical critical JJ cord-346267-l08ld2cy 260 64 community community NN cord-346267-l08ld2cy 260 65 size size NN cord-346267-l08ld2cy 260 66 and and CC cord-346267-l08ld2cy 260 67 its -PRON- PRP$ cord-346267-l08ld2cy 260 68 evolutionary evolutionary JJ cord-346267-l08ld2cy 260 69 implication implication NN cord-346267-l08ld2cy 260 70 Mitochondrial mitochondrial IN cord-346267-l08ld2cy 260 71 DNA dna NN cord-346267-l08ld2cy 260 72 sequences sequence NNS cord-346267-l08ld2cy 260 73 of of IN cord-346267-l08ld2cy 260 74 primates primate NNS cord-346267-l08ld2cy 260 75 : : : cord-346267-l08ld2cy 260 76 tempo tempo NN cord-346267-l08ld2cy 260 77 and and CC cord-346267-l08ld2cy 260 78 mode mode NN cord-346267-l08ld2cy 260 79 of of IN cord-346267-l08ld2cy 260 80 evolution evolution NN cord-346267-l08ld2cy 260 81 Effects effect NNS cord-346267-l08ld2cy 260 82 of of IN cord-346267-l08ld2cy 260 83 passage passage NN cord-346267-l08ld2cy 260 84 history history NN cord-346267-l08ld2cy 260 85 and and CC cord-346267-l08ld2cy 260 86 sampling sampling NN cord-346267-l08ld2cy 260 87 bias bias NN cord-346267-l08ld2cy 260 88 on on IN cord-346267-l08ld2cy 260 89 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 260 90 reconstruction reconstruction NN cord-346267-l08ld2cy 260 91 of of IN cord-346267-l08ld2cy 260 92 human human JJ cord-346267-l08ld2cy 260 93 influenza influenza NN cord-346267-l08ld2cy 260 94 A a DT cord-346267-l08ld2cy 260 95 evolution evolution NN cord-346267-l08ld2cy 260 96 Avian avian JJ cord-346267-l08ld2cy 260 97 influenza influenza NN cord-346267-l08ld2cy 260 98 virus virus NN cord-346267-l08ld2cy 260 99 exhibits exhibit VBZ cord-346267-l08ld2cy 260 100 rapid rapid JJ cord-346267-l08ld2cy 260 101 evolutionary evolutionary JJ cord-346267-l08ld2cy 260 102 dynamics dynamic NNS cord-346267-l08ld2cy 260 103 Hitchhiking Hitchhiking NNP cord-346267-l08ld2cy 260 104 and and CC cord-346267-l08ld2cy 260 105 the the DT cord-346267-l08ld2cy 260 106 population population NN cord-346267-l08ld2cy 260 107 genetic genetic JJ cord-346267-l08ld2cy 260 108 structure structure NN cord-346267-l08ld2cy 260 109 of of IN cord-346267-l08ld2cy 261 1 avian avian JJ cord-346267-l08ld2cy 261 2 influenza influenza NN cord-346267-l08ld2cy 261 3 virus virus NN cord-346267-l08ld2cy 261 4 Frequent frequent JJ cord-346267-l08ld2cy 261 5 toggling toggle VBG cord-346267-l08ld2cy 261 6 between between IN cord-346267-l08ld2cy 261 7 alternative alternative JJ cord-346267-l08ld2cy 261 8 amino amino NN cord-346267-l08ld2cy 261 9 acids acid NNS cord-346267-l08ld2cy 261 10 is be VBZ cord-346267-l08ld2cy 261 11 driven drive VBN cord-346267-l08ld2cy 261 12 by by IN cord-346267-l08ld2cy 261 13 selection selection NN cord-346267-l08ld2cy 261 14 in in IN cord-346267-l08ld2cy 261 15 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 261 16 A A NNP cord-346267-l08ld2cy 261 17 combined combine VBN cord-346267-l08ld2cy 261 18 empirical empirical JJ cord-346267-l08ld2cy 261 19 and and CC cord-346267-l08ld2cy 261 20 mechanistic mechanistic JJ cord-346267-l08ld2cy 261 21 codon codon NN cord-346267-l08ld2cy 261 22 model model NN cord-346267-l08ld2cy 261 23 Relaxed relaxed JJ cord-346267-l08ld2cy 261 24 phylogenetics phylogenetic NNS cord-346267-l08ld2cy 261 25 and and CC cord-346267-l08ld2cy 261 26 dating date VBG cord-346267-l08ld2cy 261 27 with with IN cord-346267-l08ld2cy 261 28 confidence confidence NN cord-346267-l08ld2cy 261 29 BEAST BEAST NNP cord-346267-l08ld2cy 261 30 : : : cord-346267-l08ld2cy 262 1 Bayesian bayesian JJ cord-346267-l08ld2cy 262 2 evolutionary evolutionary JJ cord-346267-l08ld2cy 262 3 analysis analysis NN cord-346267-l08ld2cy 262 4 by by IN cord-346267-l08ld2cy 262 5 sampling sampling NN cord-346267-l08ld2cy 262 6 trees tree NNS cord-346267-l08ld2cy 262 7 Bayesian bayesian JJ cord-346267-l08ld2cy 262 8 coalescent coalescent JJ cord-346267-l08ld2cy 262 9 inference inference NN cord-346267-l08ld2cy 262 10 of of IN cord-346267-l08ld2cy 262 11 past past JJ cord-346267-l08ld2cy 262 12 population population NN cord-346267-l08ld2cy 262 13 dynamics dynamic NNS cord-346267-l08ld2cy 262 14 from from IN cord-346267-l08ld2cy 262 15 molecular molecular JJ cord-346267-l08ld2cy 262 16 sequences sequence NNS cord-346267-l08ld2cy 262 17 Rates rate NNS cord-346267-l08ld2cy 262 18 of of IN cord-346267-l08ld2cy 262 19 evolutionary evolutionary JJ cord-346267-l08ld2cy 262 20 change change NN cord-346267-l08ld2cy 262 21 in in IN cord-346267-l08ld2cy 262 22 viruses virus NNS cord-346267-l08ld2cy 262 23 : : : cord-346267-l08ld2cy 263 1 patterns pattern NNS cord-346267-l08ld2cy 263 2 and and CC cord-346267-l08ld2cy 263 3 determinants determinant NNS cord-346267-l08ld2cy 263 4 Evolution evolution NN cord-346267-l08ld2cy 263 5 of of IN cord-346267-l08ld2cy 263 6 the the DT cord-346267-l08ld2cy 263 7 human human JJ cord-346267-l08ld2cy 263 8 immunodeficiency immunodeficiency NN cord-346267-l08ld2cy 263 9 virus virus NN cord-346267-l08ld2cy 263 10 envelope envelope NN cord-346267-l08ld2cy 263 11 gene gene NN cord-346267-l08ld2cy 263 12 is be VBZ cord-346267-l08ld2cy 263 13 dominated dominate VBN cord-346267-l08ld2cy 263 14 by by IN cord-346267-l08ld2cy 263 15 purifying purify VBG cord-346267-l08ld2cy 263 16 selection selection NN cord-346267-l08ld2cy 263 17 Paleovirology paleovirology NN cord-346267-l08ld2cy 263 18 - - HYPH cord-346267-l08ld2cy 263 19 modern modern JJ cord-346267-l08ld2cy 263 20 consequences consequence NNS cord-346267-l08ld2cy 263 21 of of IN cord-346267-l08ld2cy 263 22 ancient ancient JJ cord-346267-l08ld2cy 263 23 viruses virus NNS cord-346267-l08ld2cy 263 24 Origin Origin NNP cord-346267-l08ld2cy 263 25 of of IN cord-346267-l08ld2cy 263 26 measles measles NNP cord-346267-l08ld2cy 263 27 virus virus NN cord-346267-l08ld2cy 263 28 : : : cord-346267-l08ld2cy 263 29 divergence divergence NN cord-346267-l08ld2cy 263 30 from from IN cord-346267-l08ld2cy 263 31 rinderpest rinderpest NN cord-346267-l08ld2cy 263 32 virus virus NN cord-346267-l08ld2cy 263 33 between between IN cord-346267-l08ld2cy 263 34 the the DT cord-346267-l08ld2cy 263 35 11th 11th JJ cord-346267-l08ld2cy 263 36 and and CC cord-346267-l08ld2cy 263 37 12th 12th NN cord-346267-l08ld2cy 263 38 centuries century NNS cord-346267-l08ld2cy 264 1 A a DT cord-346267-l08ld2cy 264 2 transitional transitional JJ cord-346267-l08ld2cy 264 3 endogenous endogenous JJ cord-346267-l08ld2cy 264 4 lentivirus lentivirus NN cord-346267-l08ld2cy 264 5 from from IN cord-346267-l08ld2cy 264 6 the the DT cord-346267-l08ld2cy 264 7 genome genome NN cord-346267-l08ld2cy 264 8 of of IN cord-346267-l08ld2cy 264 9 a a DT cord-346267-l08ld2cy 264 10 basal basal NN cord-346267-l08ld2cy 264 11 primate primate NN cord-346267-l08ld2cy 264 12 and and CC cord-346267-l08ld2cy 264 13 implications implication NNS cord-346267-l08ld2cy 264 14 for for IN cord-346267-l08ld2cy 264 15 lentivirus lentivirus NN cord-346267-l08ld2cy 264 16 evolution evolution NN cord-346267-l08ld2cy 264 17 Genomic genomic JJ cord-346267-l08ld2cy 264 18 fossils fossil NNS cord-346267-l08ld2cy 264 19 calibrate calibrate VBP cord-346267-l08ld2cy 264 20 the the DT cord-346267-l08ld2cy 264 21 long long JJ cord-346267-l08ld2cy 264 22 - - HYPH cord-346267-l08ld2cy 264 23 term term NN cord-346267-l08ld2cy 264 24 evolution evolution NN cord-346267-l08ld2cy 264 25 of of IN cord-346267-l08ld2cy 264 26 hepadnaviruses hepadnavirus NNS cord-346267-l08ld2cy 265 1 A a DT cord-346267-l08ld2cy 265 2 codon codon NN cord-346267-l08ld2cy 265 3 - - HYPH cord-346267-l08ld2cy 265 4 based base VBN cord-346267-l08ld2cy 265 5 model model NN cord-346267-l08ld2cy 265 6 of of IN cord-346267-l08ld2cy 265 7 nucleotide nucleotide JJ cord-346267-l08ld2cy 265 8 substitution substitution NN cord-346267-l08ld2cy 265 9 for for IN cord-346267-l08ld2cy 265 10 protein protein NN cord-346267-l08ld2cy 265 11 - - HYPH cord-346267-l08ld2cy 265 12 coding code VBG cord-346267-l08ld2cy 265 13 DNA dna NN cord-346267-l08ld2cy 265 14 sequences sequence NNS cord-346267-l08ld2cy 265 15 Time Time NNP cord-346267-l08ld2cy 265 16 dependency dependency NN cord-346267-l08ld2cy 265 17 of of IN cord-346267-l08ld2cy 265 18 molecular molecular JJ cord-346267-l08ld2cy 265 19 rate rate NN cord-346267-l08ld2cy 265 20 estimates estimate NNS cord-346267-l08ld2cy 265 21 and and CC cord-346267-l08ld2cy 265 22 systematic systematic JJ cord-346267-l08ld2cy 265 23 overestimation overestimation NN cord-346267-l08ld2cy 265 24 of of IN cord-346267-l08ld2cy 265 25 recent recent JJ cord-346267-l08ld2cy 265 26 divergence divergence NN cord-346267-l08ld2cy 265 27 times time NNS cord-346267-l08ld2cy 265 28 Molecular Molecular NNP cord-346267-l08ld2cy 265 29 clocks clock NNS cord-346267-l08ld2cy 265 30 and and CC cord-346267-l08ld2cy 265 31 the the DT cord-346267-l08ld2cy 265 32 puzzle puzzle NN cord-346267-l08ld2cy 265 33 of of IN cord-346267-l08ld2cy 265 34 RNA RNA NNP cord-346267-l08ld2cy 265 35 virus virus NN cord-346267-l08ld2cy 265 36 origins origin VBZ cord-346267-l08ld2cy 265 37 Patterns pattern NNS cord-346267-l08ld2cy 265 38 of of IN cord-346267-l08ld2cy 265 39 intra intra JJ cord-346267-l08ld2cy 265 40 - - : cord-346267-l08ld2cy 265 41 and and CC cord-346267-l08ld2cy 265 42 interhost interhost NNP cord-346267-l08ld2cy 265 43 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 265 44 variation variation NN cord-346267-l08ld2cy 265 45 reveal reveal VBP cord-346267-l08ld2cy 265 46 strong strong JJ cord-346267-l08ld2cy 265 47 purifying purifying NN cord-346267-l08ld2cy 265 48 selection selection NN cord-346267-l08ld2cy 265 49 in in IN cord-346267-l08ld2cy 265 50 dengue dengue NN cord-346267-l08ld2cy 265 51 virus virus NN cord-346267-l08ld2cy 265 52 Evidence evidence NN cord-346267-l08ld2cy 265 53 of of IN cord-346267-l08ld2cy 265 54 the the DT cord-346267-l08ld2cy 265 55 recombinant recombinant JJ cord-346267-l08ld2cy 265 56 origin origin NN cord-346267-l08ld2cy 265 57 of of IN cord-346267-l08ld2cy 265 58 a a DT cord-346267-l08ld2cy 265 59 bat bat NN cord-346267-l08ld2cy 265 60 severe severe JJ cord-346267-l08ld2cy 265 61 acute acute JJ cord-346267-l08ld2cy 265 62 respiratory respiratory JJ cord-346267-l08ld2cy 265 63 syndrome syndrome NN cord-346267-l08ld2cy 265 64 ( ( -LRB- cord-346267-l08ld2cy 265 65 SARS)-like sars)-like CD cord-346267-l08ld2cy 265 66 coronavirus coronavirus NN cord-346267-l08ld2cy 265 67 and and CC cord-346267-l08ld2cy 265 68 its -PRON- PRP$ cord-346267-l08ld2cy 265 69 implications implication NNS cord-346267-l08ld2cy 265 70 on on IN cord-346267-l08ld2cy 265 71 the the DT cord-346267-l08ld2cy 265 72 direct direct JJ cord-346267-l08ld2cy 265 73 ancestor ancestor NN cord-346267-l08ld2cy 265 74 of of IN cord-346267-l08ld2cy 265 75 SARS SARS NNP cord-346267-l08ld2cy 265 76 coronavirus coronavirus NN cord-346267-l08ld2cy 265 77 Endogenous endogenous JJ cord-346267-l08ld2cy 265 78 non non JJ cord-346267-l08ld2cy 265 79 - - JJ cord-346267-l08ld2cy 265 80 retroviral retroviral JJ cord-346267-l08ld2cy 265 81 RNA RNA NNP cord-346267-l08ld2cy 265 82 virus virus NN cord-346267-l08ld2cy 265 83 elements element NNS cord-346267-l08ld2cy 265 84 in in IN cord-346267-l08ld2cy 265 85 mammalian mammalian JJ cord-346267-l08ld2cy 265 86 genomes genome NNS cord-346267-l08ld2cy 265 87 Evolution evolution NN cord-346267-l08ld2cy 265 88 of of IN cord-346267-l08ld2cy 265 89 protein protein NN cord-346267-l08ld2cy 265 90 molecules molecule NNS cord-346267-l08ld2cy 265 91 Endogenous endogenous JJ cord-346267-l08ld2cy 265 92 viral viral JJ cord-346267-l08ld2cy 265 93 elements element NNS cord-346267-l08ld2cy 265 94 in in IN cord-346267-l08ld2cy 265 95 animal animal NN cord-346267-l08ld2cy 265 96 genomes genome NNS cord-346267-l08ld2cy 265 97 Macroevolution macroevolution NN cord-346267-l08ld2cy 265 98 of of IN cord-346267-l08ld2cy 265 99 complex complex JJ cord-346267-l08ld2cy 265 100 retroviruses retrovirus NNS cord-346267-l08ld2cy 265 101 Adaptation adaptation NN cord-346267-l08ld2cy 265 102 to to IN cord-346267-l08ld2cy 265 103 different different JJ cord-346267-l08ld2cy 265 104 human human JJ cord-346267-l08ld2cy 265 105 populations population NNS cord-346267-l08ld2cy 265 106 by by IN cord-346267-l08ld2cy 265 107 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 265 108 revealed reveal VBN cord-346267-l08ld2cy 265 109 by by IN cord-346267-l08ld2cy 265 110 codon codon NN cord-346267-l08ld2cy 265 111 - - HYPH cord-346267-l08ld2cy 265 112 based base VBN cord-346267-l08ld2cy 265 113 analyses analysis NNS cord-346267-l08ld2cy 266 1 HyPhy HyPhy NNP cord-346267-l08ld2cy 266 2 : : : cord-346267-l08ld2cy 266 3 hypothesis hypothesis NN cord-346267-l08ld2cy 266 4 testing testing NN cord-346267-l08ld2cy 266 5 using use VBG cord-346267-l08ld2cy 266 6 phylogenies phylogeny NNS cord-346267-l08ld2cy 266 7 Site site NN cord-346267-l08ld2cy 266 8 - - HYPH cord-346267-l08ld2cy 266 9 to to IN cord-346267-l08ld2cy 266 10 - - HYPH cord-346267-l08ld2cy 266 11 site site NN cord-346267-l08ld2cy 266 12 variation variation NN cord-346267-l08ld2cy 266 13 of of IN cord-346267-l08ld2cy 266 14 synonymous synonymous JJ cord-346267-l08ld2cy 266 15 substitution substitution NN cord-346267-l08ld2cy 266 16 rates rate NNS cord-346267-l08ld2cy 266 17 Evolutionary evolutionary JJ cord-346267-l08ld2cy 266 18 fingerprinting fingerprinting NN cord-346267-l08ld2cy 266 19 of of IN cord-346267-l08ld2cy 266 20 genes gene NNS cord-346267-l08ld2cy 267 1 A a DT cord-346267-l08ld2cy 267 2 Bayesian bayesian JJ cord-346267-l08ld2cy 267 3 mixture mixture NN cord-346267-l08ld2cy 267 4 model model NN cord-346267-l08ld2cy 267 5 for for IN cord-346267-l08ld2cy 267 6 across across IN cord-346267-l08ld2cy 267 7 - - HYPH cord-346267-l08ld2cy 267 8 site site NN cord-346267-l08ld2cy 267 9 heterogeneities heterogeneity NNS cord-346267-l08ld2cy 267 10 in in IN cord-346267-l08ld2cy 267 11 the the DT cord-346267-l08ld2cy 267 12 amino amino NN cord-346267-l08ld2cy 267 13 - - HYPH cord-346267-l08ld2cy 267 14 acid acid NN cord-346267-l08ld2cy 267 15 replacement replacement NN cord-346267-l08ld2cy 267 16 process process NN cord-346267-l08ld2cy 267 17 Evidence evidence NN cord-346267-l08ld2cy 267 18 for for IN cord-346267-l08ld2cy 267 19 two two CD cord-346267-l08ld2cy 267 20 independent independent JJ cord-346267-l08ld2cy 267 21 domestications domestication NNS cord-346267-l08ld2cy 267 22 of of IN cord-346267-l08ld2cy 267 23 cattle cattle NNS cord-346267-l08ld2cy 267 24 Plagues plague NNS cord-346267-l08ld2cy 267 25 and and CC cord-346267-l08ld2cy 267 26 peoples people NNS cord-346267-l08ld2cy 268 1 Bias Bias NNP cord-346267-l08ld2cy 269 1 Due due IN cord-346267-l08ld2cy 269 2 to to IN cord-346267-l08ld2cy 269 3 Purifying Purifying NNP cord-346267-l08ld2cy 269 4 Selection Selection NNP cord-346267-l08ld2cy 269 5 · · NFP cord-346267-l08ld2cy 269 6 Two two CD cord-346267-l08ld2cy 269 7 - - HYPH cord-346267-l08ld2cy 269 8 stepping step VBG cord-346267-l08ld2cy 269 9 through through IN cord-346267-l08ld2cy 269 10 time time NN cord-346267-l08ld2cy 269 11 : : : cord-346267-l08ld2cy 269 12 mammals mammal NNS cord-346267-l08ld2cy 269 13 and and CC cord-346267-l08ld2cy 269 14 viruses virus NNS cord-346267-l08ld2cy 269 15 Measles measle NNS cord-346267-l08ld2cy 269 16 control control NN cord-346267-l08ld2cy 269 17 and and CC cord-346267-l08ld2cy 269 18 the the DT cord-346267-l08ld2cy 269 19 prospect prospect NN cord-346267-l08ld2cy 269 20 of of IN cord-346267-l08ld2cy 269 21 eradication eradication NN cord-346267-l08ld2cy 270 1 A a DT cord-346267-l08ld2cy 270 2 likelihood likelihood NN cord-346267-l08ld2cy 270 3 approach approach NN cord-346267-l08ld2cy 270 4 for for IN cord-346267-l08ld2cy 270 5 comparing compare VBG cord-346267-l08ld2cy 270 6 synonymous synonymous JJ cord-346267-l08ld2cy 270 7 and and CC cord-346267-l08ld2cy 270 8 nonsynonymous nonsynonymous JJ cord-346267-l08ld2cy 270 9 nucleotide nucleotide JJ cord-346267-l08ld2cy 270 10 substitution substitution NN cord-346267-l08ld2cy 270 11 rates rate NNS cord-346267-l08ld2cy 270 12 , , , cord-346267-l08ld2cy 270 13 with with IN cord-346267-l08ld2cy 270 14 application application NN cord-346267-l08ld2cy 270 15 to to IN cord-346267-l08ld2cy 270 16 the the DT cord-346267-l08ld2cy 270 17 chloroplast chloroplast NN cord-346267-l08ld2cy 270 18 genome genome NN cord-346267-l08ld2cy 270 19 Animal Animal NNP cord-346267-l08ld2cy 270 20 science science NN cord-346267-l08ld2cy 270 21 . . . cord-346267-l08ld2cy 271 1 rinderpest rinderpest NN cord-346267-l08ld2cy 271 2 , , , cord-346267-l08ld2cy 271 3 deadly deadly JJ cord-346267-l08ld2cy 271 4 for for IN cord-346267-l08ld2cy 271 5 cattle cattle NNS cord-346267-l08ld2cy 271 6 , , , cord-346267-l08ld2cy 271 7 joins join VBZ cord-346267-l08ld2cy 271 8 smallpox smallpox NN cord-346267-l08ld2cy 271 9 as as IN cord-346267-l08ld2cy 271 10 a a DT cord-346267-l08ld2cy 271 11 vanquished vanquish VBN cord-346267-l08ld2cy 271 12 disease disease NN cord-346267-l08ld2cy 271 13 Fauna Fauna NNP cord-346267-l08ld2cy 271 14 of of IN cord-346267-l08ld2cy 271 15 Catal Catal NNP cord-346267-l08ld2cy 271 16 Hüyük Hüyük NNP cord-346267-l08ld2cy 271 17 : : : cord-346267-l08ld2cy 271 18 evidence evidence NN cord-346267-l08ld2cy 271 19 for for IN cord-346267-l08ld2cy 271 20 early early JJ cord-346267-l08ld2cy 271 21 cattle cattle NNS cord-346267-l08ld2cy 271 22 domestication domestication NN cord-346267-l08ld2cy 271 23 in in IN cord-346267-l08ld2cy 271 24 Anatolia Anatolia NNP cord-346267-l08ld2cy 272 1 The the DT cord-346267-l08ld2cy 272 2 evolutionary evolutionary JJ cord-346267-l08ld2cy 272 3 and and CC cord-346267-l08ld2cy 272 4 epidemiological epidemiological JJ cord-346267-l08ld2cy 272 5 dynamics dynamic NNS cord-346267-l08ld2cy 272 6 of of IN cord-346267-l08ld2cy 272 7 the the DT cord-346267-l08ld2cy 272 8 paramyxoviridae paramyxoviridae JJ cord-346267-l08ld2cy 272 9 Phylogenetic phylogenetic JJ cord-346267-l08ld2cy 272 10 evidence evidence NN cord-346267-l08ld2cy 272 11 for for IN cord-346267-l08ld2cy 272 12 deleterious deleterious JJ cord-346267-l08ld2cy 272 13 mutation mutation NN cord-346267-l08ld2cy 272 14 load load NN cord-346267-l08ld2cy 272 15 in in IN cord-346267-l08ld2cy 272 16 RNA RNA NNP cord-346267-l08ld2cy 272 17 viruses virus NNS cord-346267-l08ld2cy 272 18 and and CC cord-346267-l08ld2cy 272 19 its -PRON- PRP$ cord-346267-l08ld2cy 272 20 contribution contribution NN cord-346267-l08ld2cy 272 21 to to IN cord-346267-l08ld2cy 272 22 viral viral JJ cord-346267-l08ld2cy 272 23 evolution evolution NN cord-346267-l08ld2cy 273 1 Rāzī Rāzī NNP cord-346267-l08ld2cy 273 2 ABMiZ. ABMiZ. NNP cord-346267-l08ld2cy 273 3 1848 1848 CD cord-346267-l08ld2cy 273 4 . . . cord-346267-l08ld2cy 274 1 A a DT cord-346267-l08ld2cy 274 2 treatise treatise NN cord-346267-l08ld2cy 274 3 on on IN cord-346267-l08ld2cy 274 4 the the DT cord-346267-l08ld2cy 274 5 small small JJ cord-346267-l08ld2cy 274 6 - - HYPH cord-346267-l08ld2cy 274 7 pox pox NN cord-346267-l08ld2cy 274 8 and and CC cord-346267-l08ld2cy 274 9 measles measle NNS cord-346267-l08ld2cy 275 1 The the DT cord-346267-l08ld2cy 275 2 history history NN cord-346267-l08ld2cy 275 3 of of IN cord-346267-l08ld2cy 275 4 the the DT cord-346267-l08ld2cy 275 5 acute acute JJ cord-346267-l08ld2cy 275 6 exanthemata exanthemata JJ cord-346267-l08ld2cy 275 7 Comparison comparison NN cord-346267-l08ld2cy 275 8 of of IN cord-346267-l08ld2cy 275 9 sequences sequence NNS cord-346267-l08ld2cy 275 10 of of IN cord-346267-l08ld2cy 275 11 the the DT cord-346267-l08ld2cy 275 12 H H NNP cord-346267-l08ld2cy 275 13 , , , cord-346267-l08ld2cy 275 14 F F NNP cord-346267-l08ld2cy 275 15 , , , cord-346267-l08ld2cy 275 16 and and CC cord-346267-l08ld2cy 275 17 N n CD cord-346267-l08ld2cy 275 18 coding coding NN cord-346267-l08ld2cy 275 19 genes gene NNS cord-346267-l08ld2cy 275 20 of of IN cord-346267-l08ld2cy 275 21 measles measles NN cord-346267-l08ld2cy 275 22 - - HYPH cord-346267-l08ld2cy 275 23 virus virus NN cord-346267-l08ld2cy 275 24 vaccine vaccine NN cord-346267-l08ld2cy 275 25 strains strain VBZ cord-346267-l08ld2cy 275 26 The the DT cord-346267-l08ld2cy 275 27 virion virion NN cord-346267-l08ld2cy 275 28 glycoproteins glycoprotein NNS cord-346267-l08ld2cy 275 29 of of IN cord-346267-l08ld2cy 275 30 Ebola Ebola NNP cord-346267-l08ld2cy 275 31 viruses virus NNS cord-346267-l08ld2cy 275 32 are be VBP cord-346267-l08ld2cy 275 33 encoded encode VBN cord-346267-l08ld2cy 275 34 in in IN cord-346267-l08ld2cy 275 35 two two CD cord-346267-l08ld2cy 275 36 reading read VBG cord-346267-l08ld2cy 275 37 frames frame NNS cord-346267-l08ld2cy 275 38 and and CC cord-346267-l08ld2cy 275 39 are be VBP cord-346267-l08ld2cy 275 40 expressed express VBN cord-346267-l08ld2cy 275 41 through through IN cord-346267-l08ld2cy 275 42 transcriptional transcriptional JJ cord-346267-l08ld2cy 275 43 editing edit VBG cord-346267-l08ld2cy 275 44 A a DT cord-346267-l08ld2cy 275 45 model model NN cord-346267-l08ld2cy 275 46 of of IN cord-346267-l08ld2cy 275 47 directional directional JJ cord-346267-l08ld2cy 275 48 selection selection NN cord-346267-l08ld2cy 275 49 applied apply VBN cord-346267-l08ld2cy 275 50 to to IN cord-346267-l08ld2cy 275 51 the the DT cord-346267-l08ld2cy 275 52 evolution evolution NN cord-346267-l08ld2cy 275 53 of of IN cord-346267-l08ld2cy 275 54 drug drug NN cord-346267-l08ld2cy 275 55 resistance resistance NN cord-346267-l08ld2cy 275 56 in in IN cord-346267-l08ld2cy 275 57 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 275 58 Choosing choose VBG cord-346267-l08ld2cy 275 59 appropriate appropriate JJ cord-346267-l08ld2cy 275 60 substitution substitution NN cord-346267-l08ld2cy 275 61 models model NNS cord-346267-l08ld2cy 275 62 for for IN cord-346267-l08ld2cy 275 63 the the DT cord-346267-l08ld2cy 275 64 phylogenetic phylogenetic JJ cord-346267-l08ld2cy 275 65 analysis analysis NN cord-346267-l08ld2cy 275 66 of of IN cord-346267-l08ld2cy 275 67 proteincoding proteincode VBG cord-346267-l08ld2cy 275 68 sequences sequence NNS cord-346267-l08ld2cy 275 69 Improved Improved NNP cord-346267-l08ld2cy 275 70 sampling sampling NN cord-346267-l08ld2cy 275 71 - - HYPH cord-346267-l08ld2cy 275 72 importance importance NN cord-346267-l08ld2cy 275 73 resampling resampling NN cord-346267-l08ld2cy 275 74 and and CC cord-346267-l08ld2cy 275 75 reduced reduced JJ cord-346267-l08ld2cy 275 76 bias bias NN cord-346267-l08ld2cy 275 77 importance importance NN cord-346267-l08ld2cy 275 78 sampling sample VBG cord-346267-l08ld2cy 275 79 Many many JJ cord-346267-l08ld2cy 275 80 - - HYPH cord-346267-l08ld2cy 275 81 core core NN cord-346267-l08ld2cy 275 82 algorithms algorithm NNS cord-346267-l08ld2cy 275 83 for for IN cord-346267-l08ld2cy 275 84 statistical statistical JJ cord-346267-l08ld2cy 275 85 phylogenetics phylogenetic NNS cord-346267-l08ld2cy 275 86 Model Model NNP cord-346267-l08ld2cy 275 87 selection selection NN cord-346267-l08ld2cy 275 88 in in IN cord-346267-l08ld2cy 275 89 phylogenetics phylogenetic NNS cord-346267-l08ld2cy 275 90 The the DT cord-346267-l08ld2cy 275 91 origin origin NN cord-346267-l08ld2cy 275 92 and and CC cord-346267-l08ld2cy 275 93 evolution evolution NN cord-346267-l08ld2cy 275 94 of of IN cord-346267-l08ld2cy 275 95 Ebola Ebola NNP cord-346267-l08ld2cy 275 96 and and CC cord-346267-l08ld2cy 275 97 Marburg Marburg NNP cord-346267-l08ld2cy 276 1 viruses virus NNS cord-346267-l08ld2cy 277 1 Characterization characterization NN cord-346267-l08ld2cy 277 2 of of IN cord-346267-l08ld2cy 277 3 the the DT cord-346267-l08ld2cy 277 4 1918 1918 CD cord-346267-l08ld2cy 277 5 influenza influenza NN cord-346267-l08ld2cy 277 6 virus virus NN cord-346267-l08ld2cy 277 7 polymerase polymerase NN cord-346267-l08ld2cy 277 8 genes gene NNS cord-346267-l08ld2cy 278 1 Some some DT cord-346267-l08ld2cy 278 2 probabilistic probabilistic JJ cord-346267-l08ld2cy 278 3 and and CC cord-346267-l08ld2cy 278 4 statistical statistical JJ cord-346267-l08ld2cy 278 5 problems problem NNS cord-346267-l08ld2cy 278 6 in in IN cord-346267-l08ld2cy 278 7 the the DT cord-346267-l08ld2cy 278 8 analysis analysis NN cord-346267-l08ld2cy 278 9 of of IN cord-346267-l08ld2cy 278 10 DNA dna NN cord-346267-l08ld2cy 278 11 sequences sequence NNS cord-346267-l08ld2cy 278 12 Filoviruses Filoviruses NNPS cord-346267-l08ld2cy 278 13 are be VBP cord-346267-l08ld2cy 278 14 ancient ancient JJ cord-346267-l08ld2cy 278 15 and and CC cord-346267-l08ld2cy 278 16 integrated integrate VBN cord-346267-l08ld2cy 278 17 into into IN cord-346267-l08ld2cy 278 18 mammalian mammalian JJ cord-346267-l08ld2cy 278 19 genomes genome NNS cord-346267-l08ld2cy 278 20 GP GP NNP cord-346267-l08ld2cy 278 21 mRNA mrna NN cord-346267-l08ld2cy 278 22 of of IN cord-346267-l08ld2cy 278 23 Ebola Ebola NNP cord-346267-l08ld2cy 278 24 virus virus NN cord-346267-l08ld2cy 278 25 is be VBZ cord-346267-l08ld2cy 278 26 edited edit VBN cord-346267-l08ld2cy 278 27 by by IN cord-346267-l08ld2cy 278 28 the the DT cord-346267-l08ld2cy 278 29 Ebola Ebola NNP cord-346267-l08ld2cy 278 30 virus virus NN cord-346267-l08ld2cy 278 31 polymerase polymerase NN cord-346267-l08ld2cy 278 32 and and CC cord-346267-l08ld2cy 278 33 by by IN cord-346267-l08ld2cy 278 34 T7 T7 NNP cord-346267-l08ld2cy 278 35 and and CC cord-346267-l08ld2cy 278 36 vaccinia vaccinia NN cord-346267-l08ld2cy 278 37 virus virus NN cord-346267-l08ld2cy 278 38 polymerases polymerases VBP cord-346267-l08ld2cy 278 39 Wave wave NN cord-346267-l08ld2cy 278 40 - - HYPH cord-346267-l08ld2cy 278 41 like like JJ cord-346267-l08ld2cy 278 42 spread spread NN cord-346267-l08ld2cy 278 43 of of IN cord-346267-l08ld2cy 278 44 Ebola Ebola NNP cord-346267-l08ld2cy 278 45 Zaire Zaire NNP cord-346267-l08ld2cy 279 1 The the DT cord-346267-l08ld2cy 279 2 re re NN cord-346267-l08ld2cy 279 3 - - NN cord-346267-l08ld2cy 279 4 emergence emergence NN cord-346267-l08ld2cy 279 5 of of IN cord-346267-l08ld2cy 279 6 H1N1 H1N1 NNP cord-346267-l08ld2cy 279 7 influenza influenza NN cord-346267-l08ld2cy 279 8 virus virus NN cord-346267-l08ld2cy 279 9 in in IN cord-346267-l08ld2cy 279 10 1977 1977 CD cord-346267-l08ld2cy 279 11 : : : cord-346267-l08ld2cy 279 12 a a DT cord-346267-l08ld2cy 279 13 cautionary cautionary JJ cord-346267-l08ld2cy 279 14 tale tale NN cord-346267-l08ld2cy 279 15 for for IN cord-346267-l08ld2cy 279 16 estimating estimate VBG cord-346267-l08ld2cy 279 17 divergence divergence NN cord-346267-l08ld2cy 279 18 times time NNS cord-346267-l08ld2cy 279 19 using use VBG cord-346267-l08ld2cy 279 20 biologically biologically RB cord-346267-l08ld2cy 279 21 unrealistic unrealistic JJ cord-346267-l08ld2cy 279 22 sampling sampling NN cord-346267-l08ld2cy 279 23 dates date VBZ cord-346267-l08ld2cy 279 24 A a DT cord-346267-l08ld2cy 279 25 quick quick JJ cord-346267-l08ld2cy 279 26 fuse fuse NN cord-346267-l08ld2cy 279 27 and and CC cord-346267-l08ld2cy 279 28 the the DT cord-346267-l08ld2cy 279 29 emergence emergence NN cord-346267-l08ld2cy 279 30 of of IN cord-346267-l08ld2cy 279 31 taura taura NNP cord-346267-l08ld2cy 279 32 syndrome syndrome NNP cord-346267-l08ld2cy 279 33 virus virus NN cord-346267-l08ld2cy 279 34 Dating date VBG cord-346267-l08ld2cy 279 35 the the DT cord-346267-l08ld2cy 279 36 age age NN cord-346267-l08ld2cy 279 37 of of IN cord-346267-l08ld2cy 279 38 the the DT cord-346267-l08ld2cy 279 39 SIV SIV NNP cord-346267-l08ld2cy 279 40 lineages lineage NNS cord-346267-l08ld2cy 279 41 that that WDT cord-346267-l08ld2cy 279 42 gave give VBD cord-346267-l08ld2cy 279 43 rise rise NN cord-346267-l08ld2cy 279 44 to to IN cord-346267-l08ld2cy 279 45 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 279 46 and and CC cord-346267-l08ld2cy 280 1 HIV-2 HIV-2 NNP cord-346267-l08ld2cy 280 2 Relaxed relaxed JJ cord-346267-l08ld2cy 280 3 selection selection NN cord-346267-l08ld2cy 280 4 and and CC cord-346267-l08ld2cy 280 5 the the DT cord-346267-l08ld2cy 280 6 evolution evolution NN cord-346267-l08ld2cy 280 7 of of IN cord-346267-l08ld2cy 280 8 RNA RNA NNP cord-346267-l08ld2cy 280 9 virus virus NN cord-346267-l08ld2cy 280 10 mucin mucin NN cord-346267-l08ld2cy 280 11 - - HYPH cord-346267-l08ld2cy 280 12 like like JJ cord-346267-l08ld2cy 280 13 pathogenicity pathogenicity NN cord-346267-l08ld2cy 280 14 factors factor NNS cord-346267-l08ld2cy 280 15 Immune immune JJ cord-346267-l08ld2cy 280 16 and and CC cord-346267-l08ld2cy 280 17 artificial artificial JJ cord-346267-l08ld2cy 280 18 selection selection NN cord-346267-l08ld2cy 280 19 in in IN cord-346267-l08ld2cy 280 20 the the DT cord-346267-l08ld2cy 280 21 haemagglutinin haemagglutinin NNP cord-346267-l08ld2cy 280 22 ( ( -LRB- cord-346267-l08ld2cy 280 23 H H NNP cord-346267-l08ld2cy 280 24 ) ) -RRB- cord-346267-l08ld2cy 280 25 glycoprotein glycoprotein NN cord-346267-l08ld2cy 280 26 of of IN cord-346267-l08ld2cy 280 27 measles measle NNS cord-346267-l08ld2cy 280 28 virus virus NN cord-346267-l08ld2cy 280 29 Increased increase VBD cord-346267-l08ld2cy 280 30 positive positive JJ cord-346267-l08ld2cy 280 31 selection selection NN cord-346267-l08ld2cy 280 32 pressure pressure NN cord-346267-l08ld2cy 280 33 in in IN cord-346267-l08ld2cy 280 34 persistent persistent JJ cord-346267-l08ld2cy 280 35 ( ( -LRB- cord-346267-l08ld2cy 280 36 SSPE SSPE NNP cord-346267-l08ld2cy 280 37 ) ) -RRB- cord-346267-l08ld2cy 280 38 versus versus IN cord-346267-l08ld2cy 280 39 acute acute JJ cord-346267-l08ld2cy 280 40 measles measle NNS cord-346267-l08ld2cy 280 41 virus virus NN cord-346267-l08ld2cy 280 42 infections infection NNS cord-346267-l08ld2cy 280 43 Can Can MD cord-346267-l08ld2cy 280 44 deleterious deleterious JJ cord-346267-l08ld2cy 280 45 mutations mutation NNS cord-346267-l08ld2cy 280 46 explain explain VBP cord-346267-l08ld2cy 280 47 the the DT cord-346267-l08ld2cy 280 48 time time NN cord-346267-l08ld2cy 280 49 dependency dependency NN cord-346267-l08ld2cy 280 50 of of IN cord-346267-l08ld2cy 280 51 molecular molecular JJ cord-346267-l08ld2cy 280 52 rate rate NN cord-346267-l08ld2cy 280 53 estimates estimate NNS cord-346267-l08ld2cy 280 54 ? ? . cord-346267-l08ld2cy 281 1 Point point NN cord-346267-l08ld2cy 281 2 , , , cord-346267-l08ld2cy 281 3 counterpoint counterpoint NN cord-346267-l08ld2cy 281 4 : : : cord-346267-l08ld2cy 282 1 the the DT cord-346267-l08ld2cy 282 2 evolution evolution NN cord-346267-l08ld2cy 282 3 of of IN cord-346267-l08ld2cy 282 4 pathogenic pathogenic JJ cord-346267-l08ld2cy 282 5 viruses virus NNS cord-346267-l08ld2cy 282 6 and and CC cord-346267-l08ld2cy 282 7 their -PRON- PRP$ cord-346267-l08ld2cy 282 8 human human JJ cord-346267-l08ld2cy 282 9 hosts host NNS cord-346267-l08ld2cy 282 10 Direct direct JJ cord-346267-l08ld2cy 282 11 evidence evidence NN cord-346267-l08ld2cy 282 12 of of IN cord-346267-l08ld2cy 282 13 extensive extensive JJ cord-346267-l08ld2cy 282 14 diversity diversity NN cord-346267-l08ld2cy 282 15 of of IN cord-346267-l08ld2cy 282 16 HIV-1 HIV-1 NNP cord-346267-l08ld2cy 282 17 in in IN cord-346267-l08ld2cy 282 18 Kinshasa Kinshasa NNP cord-346267-l08ld2cy 282 19 by by IN cord-346267-l08ld2cy 282 20 1960 1960 CD cord-346267-l08ld2cy 282 21 Island Island NNP cord-346267-l08ld2cy 282 22 biogeography biogeography NN cord-346267-l08ld2cy 282 23 reveals reveal VBZ cord-346267-l08ld2cy 282 24 the the DT cord-346267-l08ld2cy 282 25 deep deep JJ cord-346267-l08ld2cy 282 26 history history NN cord-346267-l08ld2cy 282 27 of of IN cord-346267-l08ld2cy 282 28 SIV SIV NNP cord-346267-l08ld2cy 282 29 Maximum Maximum NNP cord-346267-l08ld2cy 282 30 - - HYPH cord-346267-l08ld2cy 282 31 likelihood likelihood NN cord-346267-l08ld2cy 282 32 estimation estimation NN cord-346267-l08ld2cy 282 33 of of IN cord-346267-l08ld2cy 282 34 phylogeny phylogeny NN cord-346267-l08ld2cy 282 35 from from IN cord-346267-l08ld2cy 282 36 DNA dna NN cord-346267-l08ld2cy 282 37 sequences sequence NNS cord-346267-l08ld2cy 282 38 when when WRB cord-346267-l08ld2cy 282 39 substitution substitution NN cord-346267-l08ld2cy 282 40 rates rate NNS cord-346267-l08ld2cy 282 41 differ differ VBP cord-346267-l08ld2cy 282 42 over over IN cord-346267-l08ld2cy 282 43 sites site NNS cord-346267-l08ld2cy 282 44 Likelihood likelihood NN cord-346267-l08ld2cy 282 45 ratio ratio NN cord-346267-l08ld2cy 282 46 tests test NNS cord-346267-l08ld2cy 282 47 for for IN cord-346267-l08ld2cy 282 48 detecting detect VBG cord-346267-l08ld2cy 282 49 positive positive JJ cord-346267-l08ld2cy 282 50 selection selection NN cord-346267-l08ld2cy 282 51 and and CC cord-346267-l08ld2cy 282 52 application application NN cord-346267-l08ld2cy 282 53 to to IN cord-346267-l08ld2cy 282 54 primate primate JJ cord-346267-l08ld2cy 282 55 lysozyme lysozyme NNS cord-346267-l08ld2cy 282 56 evolution evolution NN cord-346267-l08ld2cy 282 57 A a DT cord-346267-l08ld2cy 282 58 new new JJ cord-346267-l08ld2cy 282 59 method method NN cord-346267-l08ld2cy 282 60 of of IN cord-346267-l08ld2cy 282 61 inference inference NN cord-346267-l08ld2cy 282 62 of of IN cord-346267-l08ld2cy 282 63 ancestral ancestral JJ cord-346267-l08ld2cy 282 64 nucleotide nucleotide NN cord-346267-l08ld2cy 282 65 and and CC cord-346267-l08ld2cy 282 66 amino amino NN cord-346267-l08ld2cy 282 67 acid acid NN cord-346267-l08ld2cy 282 68 sequences sequence NNS cord-346267-l08ld2cy 282 69 Bayesian bayesian JJ cord-346267-l08ld2cy 282 70 estimates estimate NNS cord-346267-l08ld2cy 282 71 of of IN cord-346267-l08ld2cy 282 72 the the DT cord-346267-l08ld2cy 282 73 evolutionary evolutionary JJ cord-346267-l08ld2cy 282 74 rate rate NN cord-346267-l08ld2cy 282 75 and and CC cord-346267-l08ld2cy 282 76 age age NN cord-346267-l08ld2cy 282 77 of of IN cord-346267-l08ld2cy 282 78 hepatitis hepatitis NN cord-346267-l08ld2cy 282 79 B B NNP cord-346267-l08ld2cy 282 80 virus virus NN cord-346267-l08ld2cy 282 81 Genetic genetic JJ cord-346267-l08ld2cy 282 82 variability variability NN cord-346267-l08ld2cy 282 83 and and CC cord-346267-l08ld2cy 282 84 molecular molecular JJ cord-346267-l08ld2cy 282 85 evolution evolution NN cord-346267-l08ld2cy 282 86 of of IN cord-346267-l08ld2cy 282 87 the the DT cord-346267-l08ld2cy 282 88 human human JJ cord-346267-l08ld2cy 282 89 respiratory respiratory JJ cord-346267-l08ld2cy 282 90 syncytial syncytial JJ cord-346267-l08ld2cy 282 91 virus virus NN cord-346267-l08ld2cy 282 92 subgroup subgroup NN cord-346267-l08ld2cy 282 93 B b NN cord-346267-l08ld2cy 282 94 attachment attachment NN cord-346267-l08ld2cy 282 95 G g NN cord-346267-l08ld2cy 282 96 protein protein NN cord-346267-l08ld2cy 283 1 We -PRON- PRP cord-346267-l08ld2cy 283 2 thank thank VBP cord-346267-l08ld2cy 283 3 the the DT cord-346267-l08ld2cy 283 4 Associate Associate NNP cord-346267-l08ld2cy 283 5 Editor Editor NNP cord-346267-l08ld2cy 283 6 Alexei Alexei NNP cord-346267-l08ld2cy 283 7 Drummond Drummond NNP cord-346267-l08ld2cy 283 8 , , , cord-346267-l08ld2cy 283 9 Jeff Jeff NNP cord-346267-l08ld2cy 283 10 Thorne Thorne NNP cord-346267-l08ld2cy 283 11 , , , cord-346267-l08ld2cy 283 12 and and CC cord-346267-l08ld2cy 283 13 two two CD cord-346267-l08ld2cy 283 14 anonymous anonymous JJ cord-346267-l08ld2cy 283 15 reviewers reviewer NNS cord-346267-l08ld2cy 283 16 for for IN cord-346267-l08ld2cy 283 17 their -PRON- PRP$ cord-346267-l08ld2cy 283 18 valuable valuable JJ cord-346267-l08ld2cy 283 19 comments comment NNS cord-346267-l08ld2cy 283 20 . . . cord-346267-l08ld2cy 284 1 This this DT cord-346267-l08ld2cy 284 2 research research NN cord-346267-l08ld2cy 284 3 was be VBD cord-346267-l08ld2cy 284 4 supported support VBN cord-346267-l08ld2cy 284 5 in in IN cord-346267-l08ld2cy 284 6 part part NN cord-346267-l08ld2cy 284 7 by by IN cord-346267-l08ld2cy 284 8 the the DT cord-346267-l08ld2cy 284 9 National National NNP cord-346267-l08ld2cy 284 10 Institutes Institutes NNPS cord-346267-l08ld2cy 284 11 of of IN cord-346267-l08ld2cy 284 12 Health Health NNP cord-346267-l08ld2cy 284 13 ( ( -LRB- cord-346267-l08ld2cy 284 14 _SP cord-346267-l08ld2cy 284 15 The the DT cord-346267-l08ld2cy 284 16 relative relative JJ cord-346267-l08ld2cy 284 17 importance importance NN cord-346267-l08ld2cy 284 18 of of IN cord-346267-l08ld2cy 284 19 these these DT cord-346267-l08ld2cy 284 20 factors factor NNS cord-346267-l08ld2cy 284 21 in in IN cord-346267-l08ld2cy 284 22 understanding understand VBG cord-346267-l08ld2cy 284 23 the the DT cord-346267-l08ld2cy 284 24 evolutionary evolutionary JJ cord-346267-l08ld2cy 284 25 history history NN cord-346267-l08ld2cy 284 26 of of IN cord-346267-l08ld2cy 284 27 other other JJ cord-346267-l08ld2cy 284 28 taxa taxon NNS cord-346267-l08ld2cy 284 29 remains remain VBZ cord-346267-l08ld2cy 284 30 to to TO cord-346267-l08ld2cy 284 31 be be VB cord-346267-l08ld2cy 284 32 seen see VBN cord-346267-l08ld2cy 284 33 . . . cord-346267-l08ld2cy 285 1 Supplementary supplementary JJ cord-346267-l08ld2cy 285 2 figs fig NNS cord-346267-l08ld2cy 285 3 . . . cord-346267-l08ld2cy 286 1 S1-S3 S1-S3 NNP cord-346267-l08ld2cy 286 2 are be VBP cord-346267-l08ld2cy 286 3 available available JJ cord-346267-l08ld2cy 286 4 at at IN cord-346267-l08ld2cy 286 5 Molecular Molecular NNP cord-346267-l08ld2cy 286 6 Biology Biology NNP cord-346267-l08ld2cy 286 7 and and CC cord-346267-l08ld2cy 286 8 Evolution Evolution NNP cord-346267-l08ld2cy 286 9 online online RB cord-346267-l08ld2cy 286 10 ( ( -LRB- cord-346267-l08ld2cy 286 11 http://www.mbe http://www.mbe ADD cord-346267-l08ld2cy 286 12 . . . cord-346267-l08ld2cy 287 1 oxfordjournals.org/ oxfordjournals.org/ NNP cord-346267-l08ld2cy 287 2 ) ) -RRB- cord-346267-l08ld2cy 287 3 . . .