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98/100175/99 ...- .:.:1o:.;::o/.:.:1o:.::o/.:.:1o:.::o:.:./1o:.::oC:...:H~apl'h~~70~t~%oi1l~~ei c:=====~~r;II:ee7;ubbii:aaTclattfnOJ~~ubia I Calobryales & Treubiales (leafy) L-:1i(;00nt/li'G00nt/1i'ii00O,/1i'iioo1o nana 99/100/661199 fy1onor;:lea gottschei :::;~~/53 74/100/60197 Df!?7dg~~£~p~~siiJrans .....-=-"""":"'".--:-. R/ccla SI?. 83/·/-196 100/100199/10laro/.Jf}h~i~Yli-ggpl/,~ 98/100/59/96 Asterella wallichiana Wiesnerella denudata 99/100/-/100 58/100/-199 Conocephalum conicum Marchantiopsida (Complex thalloids) ~::::::::7~~M~all;c~h~a~nt;;iaJ,p~O~/y~m;;:o,,~p~h~a_ J Cyathodium tuberosum I ~~~~=~~N~eo.t3~BfaO;j~J!Jt;~~~iS 100/100/100/100 b s8~grhg/7~~~~~J~6~O~s L..-.,."."."..".".".."."."..".".,-- caBi~~!:%tgz~sa .....-------- ~/~~~fo~!foo71~~kaynii PetalophYJlum ralfsii 100/100/100/100 Aust~?d~~~,/J6fif!J'/~~~¥ralis Fossombroma 5£' 68/100/-/60 ..... 9_9/1_0_0/9_9/1~hYllotht,~~s~/JJg67£'ia oveolata 100/100/·/89 Hattorianthus erimonus 66/-/-/88 Moerckia sp. 100/100/100/100 Symp.~9gynOPSiS filicum Moerckla 15I III 69/100/-/88 87/100/-197 100/100/100/10 Jensenia connivens 83/100/-/99 Jensenia spinosa 86/100/-/100 100/100/100/100 Pallavicinia Iyelli Symphyogyna marginata 76/100/-/- L..- ..,..._ Hymenophyton flabellatum L..- ..:.l0::::0:.:./10::::0:.:./1::::00:.:./1;;::00~ 8a~cu'aria crispula 1001100/100/100 a ¥J~lf!h~acgJf%~7a ..... ~~ p"ttf;~i~p~g~ffaata Pelliidae (Simple thalloids) L..- .;.;10;.;;;0/.;.;10;.;;;0/.:.:10;,;;;0/.:.:100~%OJtJPo~faet!a~~r;J~Yu~'h~ L .....l1!.2!00l!l/1!.2!00~/1~00~/1!.9.00~===:..::S~c~h~istO§t//;~t~~Ii(,~'blumei + Schistochilaceae (Leafy) 98/100/1001100 Jubula bogotensis 100/100/100/100 Jubulajavanica ==::-::::'lLN/~~gg~~jJ~g~~~eg~Y£.~a 77/100/86172 [§gp~6~:%t~~~r!/~OJb~gmae Acrolejeunea p'usilla Bryopteris filicina 84/-/-/- r--"1'-----_P--'y'---cnolejeunea densistipula <nnl""lT__O_mrJ!ir~~~7J7euJn~:~~~diloba 90/100/87/81 Colo!ejeune~japqnica 89/-/-/- LePto7:j:~~:: ~1f?Pt'!d~a .....-=--::-S-:'ch_iffi-:'n~~~1J?e~7~~;;,afrf;;.J1e~sale 78/100/-/- Pleuroziales, Porellales, L..------..;.;.;;;.;.;.;;.;.;.;;.;.;.;i..::..:= l[H:~pj~gg~%tf;SiS 94/100/100/100 Radula constricta 100/100/100/100 RaC:WbWJt~a~Ya~a Ptilidiales (Leafy) 96/100/55/99 81/100/56/- 100/100/100/100 Pleurozia sp. 50/-/63/- Pleurozia gigantean Ptilidium JJ15!?e°coleopsis sacculata oPtilidium pUIC~JgO?c~'JJa ulophylla 100/100/100/100 Porella vemicosa .. ~;;.;.;.;;.;.;.;;.;.;.~----- Goebellella cormgera 99/100/99/100 Lepidolaena clavigera 100/100/99/100 Gack:f,.b~ojf;~n;e~~rJg~~ri 100/100/100/100 Gackstroemia schwabei 100/100/100/100 Metacaly'pogeia cordifolia Metacalypowia a/ternifolia r~=======::.~o~ca~/~YPe:0:!g~e~ia~qu(;~;t:J~~f~usericetorum Calypogeiaceae ... 1_00_/1_00_11_00_/1_00_-e~MitW~rJ~i~CJcffgt~:grcJeJ2 .....-------~~ Mnioloma fuscum + Gongylanthus 57/100/-/- 100/100/100/100 ~c:!Yfp~~8jrc/g~ejj5u~ula 72/·/·/· I 46/100/-/61 r:;:.::=:....- Geocalyx graveolens L_~----,:""",,:~Jungermannia subulata Oo78:mciijl----- Leiocolea heterocolpos 100/100199/100 100/1001991100 Isotachis multiceps IL~~~~1-~~.Isotachis japonica 99/100/63199 Balanti°fog!1~01!~ci~~ohylla Tylimanthus laxus 90/100/971100 Marsup.idium latifolium 84/100/80/· Goebelobryum unguiculatum 9mOo78:mciQL-~~ ~__L_e_th_o_c_o_lea ~~~~~gg~j/~aruto-toganensis 100/100/100/100 Jungermannia sp. L..-__.....,,__~J...,u-n-g-ermc!;r'iffht~~i&~{r:rwoodiana '-----_~-- Gvmnomitrion concmnaPtim Marsupella emarginata -'-"'1J..----::----'- Anastrophyllum michauxii ,.-------....- Barbilophozia hatcheri 79/-/-/- Lophozia ventricosa DiploQhyllum serrulatum Jungermanniales 61/100/95/100'------r-- sCWeania !$tephanii 99/100/84/10099/100/99/1 00 8a.j1;~~i~~5/;;aoi~gurvifolia (Leafy) ~===~::::::==Cie~p~ha~/~oz~ie~~~7i~~~~ifolia L SChi~~C!l:/~:~1i~~aruensis zz§~~~J~7£1~dens 96/100/60/- ,..:.;;.;;.:.:;.;;.:.::.::.:.::.::c==--,B...a... ........==-- Acromastigum colensoanum ='J'~----:p~sil'561~§~i~/~~~~~1i~a 75/-/·/- ~Lef~~~~ijCWe1gftt:1~7a 100/100199/100 w---".__ 100/100199/100 T~Y!J~~~fgff~i(!t~s .....----..",~...,ric...,h-o-c-OlfJ7gPn~gf~~~major r!4~~?oc&~'Io~/~'/);droides 89/100/88/86 99/100195jjilODOO-0-0/1-00'-/9-9/-10-0-- P'aBh9/~hjb~t~e~e#~6~:~s 1 65/-/-/68 611-1971100 Triandrop.hYIIu/n°ftb~~~'J:J,/;eteroPhYlla ...._-- Herbenus aduncus .....----- Mastigophora diclados L..----:1:':'::00""/1:':'::00""/1:'::00""/1:'::0;;"'0-c:=-..:.:M:.!.Y/~~ff§le~rrucosa 1""""---,.;...-------------: Verdoornia succulenta 911100/100/100 Aneura pi(lgui$ L~iQQIl~~==10=0=/100=/10=0=/10:0~=3~~~~Anewa pmgUls 79/100198/100 LObati~2~%:Jif~~g~}~ Metzgeriales _l22!l22!l22!:!.22.-{52:/':/9:1I-~::::::::::::::~Riccardia L Ri[g~lJi~dj;u1f;Rg::ceavi/rea (Simple thalloids) 90/-196/99 Metzgeria lindbergii 100/100/100/100 Metzge1fa0f'[}r~~f1aeria pubescens 2/100fL8196',....------...,......,...-- Tetraphis geniculata I I [::..;;;;~~:c:::::::::~A~n~dll~e~a~e~a~ru~p~e~s~t~~/s~~'E:2~~:£!:.Sphagnum cuspidatum 0 t 100/100/100/100 Nitella translucens Takakia lepidozioides U 9rou pS Coleochaete Orbf6~t:J~/ulgaris 453 Masuzaki & al. • Systematic position ofMizutania TAXON 59 (2) • April 2010: 448-458 Fig. S. The ML tree based on Calypogeia integristipula 0 the analysis ofthe rps4 gene Calypogeia muelleriana 0 Calypogeiaceae Mnioloma fuscum 0 sequences. Numbers above the 100/100/1001100 Mnioloma fuscum 0 branches are LBP/BP/PPB/AU; 100/100/100/100 Mizutania riccardioides • .....---------............................- ... Mizutania riccardioides • in %. incubous: 0; saccubous: o 6; transverse: D; thalloid: •. o b. b. b. o o o o o b. b. 85/-1-1- b. b. b. b. b. b. b. 69/-1-1- b. b. D D A Jungermanniineae b. 99/-/100/96 b. b. b. b. b. b. b. b. b. .....----------Jackiella curvata b. Diplophyllum albicans I Barbilophozia f1oe.rkei Cephaloziineae Anastrophyllum auntum ~----- Jamesoniella rubricaulis 100/100/100/100 ~eiomyli~. anomala I Jungermanniineae (Myliaceae) Mylla taylom 69/-1-1- Lepidozia cupressina Ch!astocaulon. dendroides ILophocoleineae Pedmophyllum mterruptum Clasmatocolea vermicularis ....._- Lophocolea concreta L-~~:!.Q2L1.2£.C===:- Pachyschistochila carnosa Perssoniellineae Schistochila lehmanniana I 1-----------------------Riccardia capensis .....------- Marchantia polymorpha in the leafy liverworts by Tsubota & Deguchi (2004) might be Calypogeiaceae (Jungermanniales suborder Jungermanni ascribed to contamination by gemmae ofCalypogeia. The anal ineae) are a nearly cosmopolitan family comprising four genera: yses, however, were conducted on the basis ofthe three samples Calypogeia, Eocalypogeia, Metacalypogeia, and Mnioloma. from three different localities (Genting Highlands, Cameron Characteristic are the incubous leaves, ventral intercalary or Highlands, Fraser's Hill) in the Malay Peninsula and no trace of occasionally terminal female branches, and well-developed fragments ofplants including gemmae ofCalypogeiaceae were marsupium. Although the thalloid morphology of Mizutania detected in the samples. It is therefore highly improbable that is entirely different from the leafy morphology ofother Caly our results are derived from contamination. The present study, pogeiaceae, there are some close morphological similarities: also reveals that Pleurozia is not a sister to Metzgeriales but to (1) The rudimentary, intercalary sexual branches ofMizutania Radula (Fig. 4). Although there are gametophytic similarities are reminiscent ofthose ofCalypogeiaceae; they are short and between Pleurozia and Metzgeriales (Crandall-Stotler & aI., ofdeterminate length, and have reduced leaves and 2-3-lobed 2005), morphological differences have been observed in the bracts which are irregularly or indistinctly triseriate. Interca sporophyte: the capsule cell wall ofPleurozia is 9-10-stratose lary sexual branches seem to be a synapomorphy of Calypo and the valves lack an elaterophore (Schuster, 1984), while in geiaceae including Mizutania, but excluding Eocalypogeia. (2) Metzgeriales the capsule wall is bistratose and each valve has Oil bodies ofMizutania consist of minute granules similar to an elaterophore. It therefore seems doubtful that Pleuroziales those of many Calypogeiaceae. (3) The chromosome number and Metzgeriales are closely related. ofMizutania ofn = 9 (Inoue & Furuki, 1992) is consistent with 454 TAXON 59 (2) • April 2010: 448-458 Masuzaki & al. • Systematic position ofMizutania that of 9 or 18 in Calypogeiaceae (Tatuno, 1941). (4) Asexual stage, (2) primary stage with no underleaves, (3) juvenile stage reproduction by means of 1-3-celled ellipsoidal greenish ex characterized by constant phyllotaxis with one underleaf per ogenous gemmae on leaf margins or surface is common in each lateral leaf. Morphological variations are interpreted as the Calypogeiaceae and Mizutania (Calypogeiaceae: Schuster, the retention ofthese stages in the mature gametophyte (Grad 1969, fig. 119; Mizutania: fig. IH). (5) The massive cuticle on stein & aI., 2003). Recent molecular phylogenetic study re the thallus surface of Mizutania is reminiscent of the cuticle vealed that Cololejeunea metzgeriopsis is not phylogenetically found in several species ofMnioloma (Mizutania: fig. lC-F, H isolated from other Cololejeunea species (Gradstein & aI, 2006; and K; Mnioloma: Schuster, 2000, figs. 206, 209, 210, 21OA). Gradstein & Wilson, 2009). Protonemal neoteny is usually In the phylogenetic tree based on rbcL sequences (Fig. 4), interpreted as an adaption to the growth in short-lived habitats Calypogeia, Metacalypogeia, Eocalypogeia, Mizutania (Ca such as on living leaves where rapid maturation and completion lypogeiaceae), and Gongylanthus (Arnelliaceae)