Symphyodon leiocarpus, sp. nov. (Symphyodontaceae, Musci) from Thailand, Classified in the New Subgenus Macrothamniopsis

2009·1.9 MB·English

by  Akiyama, Hiroyuki

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ISSN1346-7565 ActaPhytotax.Gcobot.60(2)8:7-96 (2009)

Sympllyodon leiocarpus, (SymphyodontaceaMeu,sci)
sp. nov.
from Thailand,Classifiedinthe New Subgenus Macrothamniopsis

HIRoyuKI AKIyAMAI' HIRoMI TsuBoTA2
AND
i Mi ise um o f N b t u r ean dH um an Ac ti vi ti es . f i > ,o g o ,l dy o ig a o'kMa)-'6y, oSjainmdaa-shi, tlyogo 449-1546, .h4?an,
"aki,yama@hitohaku,ip lautherf ocorrresnondencel; ' IVZitara Blotanical Carden,
Graduate Sehool ofScienc eHi,roshima Uhivenyit Jy1,56-2 Mitsumaruko-yama ,Miyojima-cho,
llatsukaichi- sHhiir,oshima Z39-0543, Jopan
Symplp?od blneiecarpu ss,p, nov., previously reported under the name Symphyodb noblongijbiius, is de-
scribed from Thai]an calnd Myanmar. The longe raxillary hairs u,pright capsules with smooth exotheciaL
cells, superficial stomata, and large rspore size suggest it sremote relationship to other members ofthe
genus. Based on a molecular phylogenctic analysis using sequences ofchloroplast rbcL gene, a new sub-
genus, S),nupbyo dsubbgn. Mticrothamniopsis, is established to accommodate the species, Morphologi-
cal comparisons with related genera ,especially those ofthe Hylecomiaceae and Symphyodontaceae ,are
also discusse dbriefly.
Key words: rbcL, S?mpityod olneiocarpus, 5),mphyod osnubg, Mtxerothamniopsis, Symphyodontaceae,
Thailand
Doi (Mt) Inthanon, the higbes tpeak in Thai- bryophyte element, contains a high diversit yof
land ,has long been known to support one of the species, which characterizes the fior aof Doi In-
richest bryophyte flora sin Asia (Dixo n1932, thanon, Nine species of 5Iynephyodb nhaye been
1935, Horikawa & Ando 1964, Kitagawa 1967a, reported from Chiang Mai and six of them are
b, Noguchi i972a, 1973, Ogawa et al. 1961, Tbuw from the upper e]evations of Doi Inthanon (He &
1968). The upper elevations are covered by well- Snider2000).
forestsI.n One (Akiyamaj, from
preserved evergreen rnontane addition, ofus with permissien
mossy forest sare developed around the summit the Royal Forest Department of Thaitand ,was
and in places over 1800 m (Hara 2003). abie to study the bryophyte fiora at the upper el-
Among the bryophyte sknown from Doi In- ev ations of Doi Inthanon in December 20eO. Lat-
thanon are a number of species endemic to the er in 2008 (Marc hand DecembeD, Akiyama
mountain and species with severely restricted again visited Doi Inthanon as a rnember efa co-
distributio nss,uch as Curvicladium kur:ziioperatiye research preject between Kyoto Uni-
(Kindb .E)nroth, Dixonia orientalis (Mitt .H), versity and Maejo University .Parts of the results
Akiyama & Tsubota ,.Nbguchiodlenduon sphaero- based on the coliections have been published in
carpum (Nog. N)inh & P6cs, Penzigietla coxdota $eparate papers (Ak'iya rn20a07, Akiyama &
(Harv,M.) Fleisch.,PinnateUa En- Tsubota 2001,2004),
amb(zpbylla
roth, and Rgeudotrismegistia undutata (Brot h&. Among the collections. we noticed a curious
Ylisuda jH. Akiyama & Tsubota .Among these, moss with the gametophytic feature osf SvmpIu7-
SyJuphyodb na ,representative of the Himalayan oclon, but lackin gthe characteristic echinate or
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spinose capsules, Superfici astlomata on the neck have been deposited b,ut lik eHe & Snider (1992,
ofthe capsules and large rspores also separated it 2000) we were unable to locat eit A.lthough the
from other species of SympItyod (oFnig .1) .The identity of S. oblongij?)lius cannot be know with
plants w,hich we describ ebelow as Svmphyodon.certainty in the absence of a type specimen, we
teiocarpus, were abundant on the base of tree attempted to characterize it from the original de-
trunks and on the branches of shrubs in the un- scription (Renau &l dCardot 1896) and from an
dergrowth along the Summit and Angka trails additional descripti obny Gangulee (1977 w)ho
around the summit area as well as along the Kew claimed to have checked the type specimen, Ac-
Mae Pan Nature r[hrail. From our fiel dobserva- cordingly, Syrupl4io doboionngij?)lius can be
tions, the new species is not rare in the local characterized as fbllows :1) shoots irregularly
mossy fbrest sS.ympPtyod ioeniocarpu swas also plnnatel ybranched ,2) leayes elliptic-oblong,
found growing on branches oftall, emergent trees concave, rounded but gradually pointed to the
in the upper montane fores tat 1600-1700 m alti- apex, 3) laminal cells linear m,ore or less prorate
tude. After ehecking previou sreports on the at upper corners, and 4) costa very short. These
moss flora of the mountain, it became clear that featur efsi twell with those of typical species of
th)mphyoc lleoinocarpu shad previously been re- Symp1tyoc lanod n i,t seerns reasonable that Broth-
ported under the name ofSympbyodon oblongijb- erus (i907 t)ransferred the species firo mLeptohy-
iius (Renau l&d Cardot) Broth. [Basiony Lmq:p- menium to Symphyodb nAt.though the gameto-
tohymenium obtongij?)Uum Renautd & Cardot. phytes are not divers eamong the species of
TYpe: India orjentalis, Sikkim, Daijeeling, L. Ste- Elymplryodb annd gametophyti cfeature asre often
vens s,n.] (Touw )968, He & Snider 1992, 200Q). scarcely useful to distingui sihndividua slpecies,
"fouw
(196 8r)eported S. oblongijblius frem the bympJryod eleniocarpz ahsas the fbllowing
four localiti eisn Chiang Mai County including feature s(Fi g1.) not observed in S, oblongijfolius:
Doi Inthanon (se eparatype sbelow) .Later, Si He 1) shoots mDre or less regularly bi- or tripinnately
(200 9i,nterne rtesource) cited three specimens of branched ,2) leave sovate-lanceolate, concave,
Tbuw's collection as S, obiongij?)lius, but did not obtuse or broadly acute, 3) laminal cells linear,
refer to others collected from Doi Inthanon. He & evenly thick walled, and totally smooth, and 4)
Snider (1992 2,000) excluded the species in their costa often reaching half of leaf length ,more or
revision of the genus, mainly because they were less fbrked. We therefore conclude that Symplry-
not able to locat ethe type specimen of S. oblon- oclon oblongijbgius and the plant swe name S.
gij?)ii uisn their search through specimens in BM, reiocaicp urespresent two differe nstpecies.
NY and other herbaria H.e & Snider (199 220,00)
therefore lef tthe taxonomic treatment for future Comparison with other species ofSyrnphyodon
investigation S.ince that time the species has not According to detaile gdeneri cdescriptio nons
been fult yreexamined. Synrphyod bbyn He & Snider (2000 )and our own
The main purpose ofthis paper is to settle the ebservations, Svnrphyod loeniocarpu sand other
identit yof Eb?mplyod ioeniocarpu sthrough mQT= members of the genus share a number of features
by fo11ows:Plantsdioicous;
phological comparisons and molecular phylo- as central strands of
geneti canalysis with related genera. stems absent or weakly devetoped; alar regions of
branch leaves differentiated
stem and slightly
MorphologicalInvestigations
with a small numbers of quadrat ecells; eostae
short and double ;perichaetia terminal on short
Comparison with Syrnphyodon oblongifolius branches and becoming extensively elongate af
We attempted to find the type specimen of ter fertilizat itoon form distincti vveaginulae
Sympbyodo onblongijZ,lius by requesting the help (Fig .1 c & d); upper portion sofsetae distinctly
of curators and keepers ofthe herbaria BM, H, L, mammillose; capsules erect and ovoid with long
NY and PC, in which Renauld's collections might beaked opercula; peristome softhe IVkckera-type
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and exostome teeth lack dorsa lmedian furrows; them, Chaetomitriopsi Csh,aetomitrium and Uit-
endostome reduced, with low basal membranes ctojackia resemble each other in their gameto-
and linear segments (sometim aebssenO and lack- phytic feature s,while Symp1tyod olneiocaflpus
ing ciiia. shares only the fbllowing features with them; (1)
SJmpbyodbnleiocarpus,however, differs (2) differentiated
reduced endostome, weakly
from the other species of Syi?u]lty iond tohen fbl- alar reg{ons, and (3 )elongated vaginula afier fer-
]owings (charac sttaeters observed enly in the lat- tilizatio (nIgnat eot val. 1999), As fbr 7}'acdythe-
ter are shown in square brackets )1:) laminal cells eium, it spendent capsules with a perfect I()/p-
of stem and branch leave ssmooth [more or less num-type peristom eand mammillose exothec{al
prorate or prorulose, often spinose, at upper cells might indicate a remote relationship to Sym-
ends], 2) axillary hairs uniseriate, (4 or ) 5-celleds pdyoclon leiocaipu sA.ccording to our field ob-
upper (2 oD 3 cclks elongate, hyalin eand basa l2 servations on the Borneo lsland ,the monospecii
celts quadrate, brownish [browni sorh hyaline, ic genus Dimorphoclaclon is unique in livin gon
uniseriate, 2 or trare l3Y-c)elled ,with a single, the leave sof shrubs growing beside streams and
quadrate, basa lcell], 3) capsule walls smooth rivers, Its upright, more or les sfi at branches with
[echin aort espinose], 4) stomata superficia] [im- attenuate tips bearing filamentou sgemmae in the
mersed], and 5) spores 20-25 stm in diameter leaf axi{s give Dimorlphoclado nan appearance
[12-2 0pem in diametetl T.hese feature sw,e cen- similar to species of Ctastobf:yopsi sM, Fleisch,
sider, clearly suggest 51ympdyodon leiocarpus to [Akiyam a1993 (a sAptyche Uasp,), Akiyama
be remotely related te other members ofthe genus 1997]. Although Dixon (1922 p)ointed out the af
Sympiryod oInn .addition, we noticed that bym- finit ybetween DimoT]phoeladb nand Chaetomi-
pbyodlon leiocap:pu sshows a diflft)r ehnabtitat trium, we do not eonsider the long, smooth setae,
preferenc e.The other species of SympJny?odocnylindrical capsules, long-beake dopercula, and
grow on the branches of undergrowth shrubs in filamentou sgemmae of DimoF;phoclado nto sug-
well preserve dfbrest sw,hile Symp1tyod loenio- gest a close relatienship. As a result, Sypaphyodon
carpus is restricted to sunnier places ,such as the leiocarpus shares most morphological features
base of tree trunks and on the branches ofshrubs with other species of Symplryodon.
at the edge of thickets and on the higher branches
oftall trees. These difilerenc aeres not sufficient, Comparison with otherfamilies
however, to classify Sympbyodo lneiocarpu sin a Upright capsules with rnore er less reduced
separate genus because species with widely dif endostornes, leaves with obtuse or widely acute
feren tfeature soften form monophyletic groups apex and often bifi dcosta as well as the epiphytic
and autoapomorphies cannot be used to infer habita tof Sympla,od loeniocarpu ssuggest affini-
phylogeneti caffinity, For example, species with ty to member of the Entodontaceae, especial.ly to
echinate and smooth capsules sometimes belong Entodon MUII, Hal, Sympbyodb nleiocaepus,
to the same genus (e. gi.n Chaetomitrium ,Aki- hewever, does not have features fbund in Ent-
yama & Suleiman 2001). odon, such as the wide alar regions with numer-
ous, small, quadrat ecells, cylindrica} capsules,
Comparison with other genera in the Sympby- and much reduced, fragmented endostome often
odbntaceae adhering te the inner surface o'f exostome teeth.
Based on morphological feature sand molecu- Sympbyodo lneiocarpu salso appears similar
lar analyses, Goffine tet al, (2008 l)iste dfive gen- to some me'mbers ef the Hylocomicaceae, espe-
era in the Symphyodentaceae, Along with Sym- cially to those with bi- or tripinnatel ybranched
pdyodbn, they are Chaetomitriops iMs, Fleisch,, stems and!or upright capsules, which can be sum-
Chaetomitrium Dozy & Molk., Dimorphocladon marized as fo11ows:
Dixon, 7>"achytheci uMm. Fleisch .a,nd Uhcle- Leptobymenium Schwagr, Although the Asian
j'acki aIgnatov, T.J. Kop. & D.H. Norris, Among representative of Leptoh.ymenium. L, tenue
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s
,
if.
u
k
FiG. 1. Stvmph>todon leiocarp uHs. Akiyama & Tsubota. a: p)ants innatura] habitat .b: dried female plant swltb mature sporo-
phytes. c: female plant with elongated vaginula inwet condition. d: elongated vaginula after fertilizu'ti (onarro iwndicatcs
unferti]lzcd peri chaetjum). e & f: branch leaves .g: stem leaE h: apex ofbranch leat i: upper margin o± branch lea£ j :me-
dian larnina lcclLs ofbranch lea£ k: alar region of branch leat 1: capsute neck and mamrnillose, uppermost portion ef a
bcaked
seta. m: amature eapsule w{th oblique opcrculum, n: closeup ofdried pcristome .o: superficial stomata at neek ef
capsu]c. p: spores. q; outeT surfacc of 1aneeo]at ecxostome teeth and filamentous endostome segments. Scales == 5 mrn in
a, b & c, 1 mm in d, O.35 mm in e-g. O.5 rnm in m & ll, and 50 ptm in h-k, o-q,
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(Hook .S)chwagr, ,has upright capsules, they are same localit oyn Doi Inthanon ,along with an ad-
not ovoid but long cylindrical. Additionall yth,e ditiona slarnple ofS. scaber fror nMyanmar.
(pro-peristomier)regu- Leptobymenium (Svmphyodon
presence of prostome as oblongij?)lius
lar deposit son the outer surfaee of the exostome was once classified therej and related genera of
teeth and well-differentiated decurren talar re- the Hylocomiaceae as well as species of Entodon
gjons also suggest a remote relationship, were also included in the analysis because they
Leptocladiella M. Fleisch ,Rather deli acte and show morphological similarities to Sympityodon,
simple shoots with few later albranches give a Other OTUs are detaile din the Appendix.
unique appearance to Lqptocladiegla among the Investigatio nof topology candidates were
Hylocomiaceae. While th,mpbyod bieniocarpus performed with the fo11owing programs: MEGA
is more or les sbi- or tri-pinnatel ybranched .Ac- 4 (Tamur eat at. 200Z Kumar et aL 2008) ,Mol-
cording to Rohrer (1995 t)h,e costa of the branch phy 2.3b3 (Adac h&i Hasegawa 1996), MrBayes
leaves is often single and ends in a spine and the 3,i.2 (Huelsenb e&c Rkonquist 2001, Ronquist &
outer surfaces of exostome teeth are basali ystri- Huelsenbeck 2003) ,PAUP" 4.0blO (SwofTbrd
inLeptocladiellapsilu(rMaittM.,)Fleisch. 2003),PAUPRat (Sike&s Lewis 2001)
ate over
Macrothamnium M. Fleisch. A number of PAUP, PhyML 2.4,4 (Guindo &n Gascuel 2003).
species have been reported under Macrothamni- Kakusan3 (Tanab 2e007) was used to select the
um. Most of them are eharacterized by having nucleotide substitution model fbr the data set. Re-
more or less sub-arcuate capsules, and finely stri- calculation of likelihood values for each tree to-
ate exostome teeth .The sele ex ¢eption is Ml lep- pology was perfbrmed with PAUP. The candidate
todymenioides Neg., which has upright and sub- topologies were evaluated by the approximateun-
globose capsules (Noguch i1972b), The long, biased test (AU; Shimodaira 2002, 2004) using
double costa, distinct ldyentat eleaf margins, CONSEL (Shimodai &r aHasegawa 2001) .Pro-
shortly beaked operculum and the basall yfinely cedures of sequencings, alignments, and phylo-
striate exosteme teeth, however, are quit edifferL geneti canalysis adopted are nearly the same as
ent from those found in SvnzF)hyo adndon in S, detaile din Tsubota et aL (200 42,005) and Ozeki
teiocarpus. ' (2007).
et al.
Orontobryum Mitt, ex M. Fteisch. resembles The phylogeny of Synzphyoclo annd it sallies is
Symphyodo lneiocarpu sin the upright, globose shown in Fig. 2 based on the alignment of 1,220
capsu]es. The paraphytlia on the stems and bps of the 42 moss chlQroplast rbcL gene se-
branches ,monopodial branching pattern ,strong quences, depicte dby the best-support etdree with
dentit'io onfthe leafmargins, and dome shaped high ranking log-likeliho ovadlues that passed
opercula with scarcely elongate beaks (Rohrerthe approximate unbiased test (AU ;Shimodaira
1985) ,however, may indicat ea remote relation- 2000, 2002). Four samples trhr espeecies) ofSym-
ship to both th?mpdyoc lino gneneral and to S. pPryodbn formed a monephyletic clade with S.
Ieiocarpu sin particular, teioeai:pu sappearing as sister to the clade with
high (AU7LBPXBPmpfBPnj/
supporting values
Molecular Phylogenetic Analysis and BPml!PP ` 1001IOO199flOOflOOfl .OabObr;evia-
Its Implications tions of these values are explained in the legend
for Fig, 2). The results support our supposition
Tb clarify the phylogeneti cpesitio nof Sym- based on morphological comparisons that S. geio-
plp,odon leiocarpus ,we carried out molecular carpus is heterogeneou samong members of th?m-
phylogenetic analyses based on chloroplast rbcL phyodon.
gene sequences (Fi g2. and Appendix) .We select- Since we were not able to includ eany samples
ed three species of Sympltyod bonne, each from from the Symphyodontaceae except for bymphy-
the three infragener igcroups recognized by He odon, it is still ambiguous whether Symp1ryodon
& Snider (1992 T)h,ey were collected from the leiocarpt tshsould be treated in a genus separate
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AUfLBPIBPmptBPnjlBPmltPP
85
--ISES.i.i.r
'blyVocorniurnspiendensAB024662
.sfigP.SIewpg.
ft.oo
LoeskeobryufncavilbliunTAB02465B

rio uJ7GfG7ra99 FlytocomiastrumpyrenaicumAB024660
he"roziumschreberiAB024664
'r an-Jd.41f.:t'it9 7b me n Cy p nu m ni t en sSAaBnei2o4n6i7a 6uncinata AB332276
ylSS7 CretoneuronfiiicinumAB0952TO
CampyfepltytilumhaifetiAB332270
.153S./.1.Y. ioorgstgewggiggii,oo EhlodonconcinnusAB332277
67
B039675
97J541.
t u
O.Otsubstitutienslsite
FiG, 2. Phylogeny of Symplo,od abndn it sallies based on l220 aLigned bps ofthc 42 rnoss chEoroplast rbclL genc sequ ¢nces,
depictedbythebest-supportedtree(GTR+Gmodel;lnL=-59lO.4)witbhighestlog-likelihoodvaluethatpassedtbeap-
proximate unbiased test (AU; Shiinodai r2a0e2, 20e4) by CONSEL. The root is arbitrarily pluced on the branch lead{ng
to PlagiomniumJ'aponicwm. The horizont atiength of each branch is propDrtiona lto the estimated number ofnuclcotide
substitutions, Supperting values greater than 50% (AUILBPZBI)nrpfBPrij/B aPrem ]shIovPvnP) near the nodes: the AU
probabiliti ebsased oll AU test using the multiscale boetstrap technique {AUs in %); IQcal bootstrap probabiliti ecsalcu-
late dwith thc estimation using the RELL resampling ]nethod (Kishi neto al. 199O, Hasegawa & Kishino l994) by Molphy
2,3b3 with TN93 model (LBPs ;in %); classical bootstrap probabiliti e(sFelsens t1c98j5n) bascd on 10,OOO replications
calculated wnh MP and Nj mcthods by MEGA 4 with maximum composite likelihoo dmodel (BPmp sand BPnis; in %);
and bootstrap probabiliti ebsased on 1,OOO replications calculated with ML methDd by PhyML with GTR + G model
(BPmls i;n %); und the Bayesian postcrio rprobabilit ycaleulated by thc Bayesian informarion criterion (BIC) approxirna-
tion (Schwar 1z978, Hasegawa & Kishino 1989) by MrBuyes with GTR + G model (PPs).
fror nSympdyocl oor nnot. Relativel syhort branch- ALfl7niti etso the Entodbninceae and llylocomia-
es among the five samples, if compared to other ceae hased on moleculdr analysis
congeneric cases in Entodbn and Plagiothecium The tree shows a relationship between Sym-
in the present analysis, however, might support pdyoclon (includ {S.n Igeioearpzas )and Entodon,
the latte prlacement. although the supporting values are low. Because
of insufficie tnatxon sampling their relationships
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are unclear; some topologies with high likelihood age; paraphyllia and pseudoparaphyllia absent.
values showed a close relationship between Ent- Rhizoic tresstricted to ventral surface of prostrate
oclon and other genera and families ,such as shoots, tufted ,smooth, brown, Stei nleave sovate-
T:huidium and the Sematophytlaceae, as shown lanceolate ,ca, 1 rnm long ,obtuse or broadly
by Tsubota et al. (2004), acute, upper surface often concave, neither pti-
None of the members of the Hylocomiaceae cate nor rugose; margins minutely crenulate; cos-
CEI[yloco milluymto,comiastrum, Leptoilymeni- ta double or more or les sbranched ,reaching ]!3-
LeptocladiellLao,eskeobF:yum,Macrotham- 1f2leaflength, laminal
um, rarely obscure; uppermost
nium, and Pleurozium )is closely related to the cells short!y rhomboid, evenly thick walled,
(Symplo,o d+o Sn. Ieiocarpus c)lade in the pres- smooth; median laminal cells linear-rhomboid,
ent analysis as was already shown by Goffine tet evenly thick walled, 40-60 ptm long ,smoothl
al. (2008 I)n. addition, these seven genera are re- marginal laminal cells becoming narrower,
motely separated into three diifere ntclades, smooth; alar region differentiate wdith small sub-
which may suggest paraphyly. quadrat ecells. Axillar .hyairs hyaline u,niseriate,
Taking into consideration the evidence dis- (4 oD 5-celle du;pper (2 oD 3 cells elongate, hya-
cussed above, we conclude that the th,nrpbyodb1n'in es,moothl basal 2 cells quadrate ,brownish,
leiocaszp ushsould be treated as new and that it Branch leave ssimilar to stem leaves if lshape and
represents a new subgenus within Svmphyodon. size; terminal branch leave smuch smaller than
stem and branch leaves ,shortly ovate, ca. O.5-O.7
Taxonomic Treatment long, deeply broadly
mm upper surface concave,
lam-
ebtuse, often minutely apiculate; uppermost
Symphyodon subg. Macrothamniopsis H. Aki- inal cells shortly rhomboid, median laminal cells
yama & Tsubota ,subg. nov 30-60 ftm long ,smooth; margins minutely cren-
Aranis Synrp]tyod osnubtgi, Sympbyodont 4sed capsul{s ulate; alar cells weakly differentiat Deido.icous:
laeyibu ssternatibus superfieialibus sporis majoribus, 20- male plant sslightly more slender than female
25 ti tn diamet ro diveTsus. plants P.erigonia on ascending shoots (usua olnly
7)pus. Symplu;o dlbeniocarpu Hs. Akiyaina & Tsu- secondary and terminal branches) ,containing
bota
3-5 antheridia. Perigonial leaves shortly ovate,
concave, to O.6 mm long, ecostate. Perichaetia
Symphyodon le;oearpus H. Akiyama & Tsubo-
sessile, restricted to ascending stems and second-
ta, sp. noy. (Fi g1.a-q) ary branches. Perichaetial leaves graduall ynar-
Affini s,SJmphyodonti erinaceo, sed capsutis laevibu sdi- rowed from ovate to oblong sheathing base, ecos-
versus. tate ,laginul amuch elongated after fertilization
n,pus .THAILAND, Chiang Mai, Doi Inthanon, and becoming a short distin cbtranch .CaL}tptrae
a2r0o0un0d,H tLheA kslumymaimta, 7mo:nlumze-nt51(ho tlroaBiK-lF, ;255i0s om- BalMt,, DCecM.U ,5, unknown. Setae reddish brown, 5-6 mm long ,al-

(rarely
HIRO, HYO, L, MO,NY, SING). most always straight slightly curved
downward distal jusbtelow
at end), mammillate
Plant sgreen or olive-green, older part sturn- neck. Capsules globose or shortly ovoid, upright,
ing yellow ,2-3 cm long, menopodially branched, surface smooth, reddish brown, slightly glossy;
prostrate ,with a number of short ascending exothecial cells hexagonal ,evenly thick walled;
shoots, forming dense mats on substrata, sub- annulus differentiate din 2 or 3 rows; stomata
dendroid ,more or les sregularly bi- or tripinnate- 6-15 per capsule, superficial, restricted to neck.
ly branched; branches terete or sometimes slight- (Jiperctt loabliquety narrowly rostrate. Peristome
ly fbliate, O,5-1.e long; double, in
complanately usually cm opening outwards rnoist conditions.
central strands absent or weakly developed ;cor- E)costome teeth 16, brownish yellow ,apically pale
tical cells lax ,thin walled, epidermal cells thick white, narrowly lanceolat en,iedian furrow ab-
walled, with substereids, tinged reddish brown in sent, 400-500 ptm long ,finel ypapilles tehrough-
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out on both surfaces. Endostome with low basal and H. Deguchi (HIRO) for searching for the holotyp eof
mernbrane, segments linea rl,acking perfbrations, Leptohymenium oblongijZ)lius. We also thank the Ana]y-
2E0P0o-3r0e0s ptm long, finely pafpiilnleolsye lcilia abse2n0t.- siin s sCeqeuenntciengr ofLif eand S ctihee nNeaet H,uirrao lsShciimean Ucn eiCveenrtseir tf ybf,r oBr haestipc
spherical, greenish, papillose , Research and Deve]opment, Hiroshitna University, for
25 Iim in diameter. supplying cryogen, Dr, W. R Buck, The New YOrk Bo-
Distribution .Myanmar (Chi nState ),Thai- tanical Garden, and Dr. D. BeufTbrd ,Harvard University
land(Chiang-MCao.i), Herbaria ,for correcting the Englis htext ,and two aneny-

Hbbitat,Forming base mous referees fbf their valuable suggestions on the draft.
small patches en of This study was financiall ysupported by Grants-in-Aid
tree trunks and on branches of shrubs and small firem the Japanes eMinistry ofEducation, Culture S,ports.
trees (sometim epesndulous )in mossy or eveF Scienc eand fechnology (No.18570 0t9o 2th)e senior au-
green upper montane fbrest sO.ften i'ntermingled thor, (No,090411 6an5d 13375003) to Dr. J, Murata (rlb-
with Macrothamnium submacrocarpum and oth- kye University) and (No ,20405008) to Dr. M. Kanzaki
(KyotoUniversity),
SvmpIo,oclon.
er species of
Distinguishing.featuT1e)s.leaves
ovate-lan-
References
ceolate, concave, obtuse or brQadly acuminate
double ]essbranched 2)
with or more or costa,
alar regions weakly developed with quadrate Adachi, J. & M. Hasegawa, 1996. MOLPHY version 2.3:
cells, 3) laminal cells linear e,venly thick watled, programs for molecular phylogeneti ebsased on max-
and totall ysmooth, 4) axillary hairs uniseriate i1m-u1m50 l.ikelihoo Cdo.mputer Science Menographs 28:

with a single basal cell, 5) capsules upright, to- Akiyama, H. 1993. Taxonomic studies of mosses of Se-
tal ly smooth, 6) stom ata superficial , and 7) spores ram and Arnbon (Molucca sE,ast Malesial collected
20-25 psm in diameter, by Indenesian-Japanese botaniea lexped{tions. VII.
Lembophyllaceae, Splachnaceae ,Myuriaceae ,Nec-
Paratypes, THAILAND. Chiang Mai: Doi Inthanon, keraceae, Brachytheciaceae Amblystegiaceae and
around the summit, monument trail ,2550 m alt, fi :Aki- SematophyLlacea ewi,th additions to previou sreports
yama Th-127 & 71i.-12 (8HYO!) ,A. 7b"w 9780 (Ll) i;bid., on Dicranacea eand Pterobryacea eJ.. Fac. Sci .Univ.
A(nHgY OK!a) ,Htr/aiAlk, l25ySaOm am2 1al5L5, 1l&l 2Afk5ly5a2m(aH Y7O1h!-)3i4 ;b&i d7.1K,Je}w-42 Tokyo, Bot. 15: 2r1f9a-x2on5o4m.ic
Akiyama, H, 1997, studies of mosses of Se-
Mae Pan nature trail ,2200 m alt., H/ Aklyama 2I576 ram and Ambon (Molucca sEa,st Malesia) cotlected
(HYO!) ;ibid. 2300 m alt, A. 7buw JO128 (L! )i;bid. by Indonesian-Japane Bsoetanical Expeditien sVIII.
2150-2250 m alt., A. 7buw 10250 (IJ ;ib{d. ,21.50-2200 m Meteoriaceae ,Hookeriaceae ,and Trachypodaceae.
alt., A. Tbuw IO065 ,JO068 & 10077 (L! )2;000 m alt., A. Human and Nature 2: 9-31,
7buiv 10010 (L! )ib;id. ,1600-l650 m ait., ll ZAklyama, Al Akiyama, H. 2006. New reeords of rr]osses from Thai-
Kanzaki, T frie & HL Ando 314 & 332 (BKF! ,CMU!, land. Trep. Bryol, 28: 59.
HYO!); Doi Suthep ,summit ofDoi Pui, 1600-1650 m alt., Akiyama, H. & M, Suleiman, 2001. TaxonomicaL notes on
A. 7bu,v 8539 (L!). the genus Chaetomitriu m(Hookeriac Meuascei,) ot'
MYANMAR. Chin State :Natma-Tftung National Borneo. Hikobia l3: 491-509.
Park, Mt, Vietoria, 2470 m alt., X n4iira teat aZ, 23262 Akiyama, H, & H. Tsubota. 2001. Rseudotrismegistia H.
(HYO!).
Akiy, & Tsubota, a new genus fbr the Semateptyylla-
2Vbte .Tbuw (l96 8al)so reported Slympdyoclon Ak i ycaemaea.. A Hct. a& P Hhy, tTostuabxo.t Gae.o 2b0ot0.4 . 5M2o: r8p5h-o-l･9o5g.ica land ge-
leiocarpusfrom Doi Chiang Dao, Chiang Mai
netical variations and familial affinity of the genus
Co., Thailand (A .7buvv 9380, L), but we have net Dixenia (Musci )Br.yol. Res. 8: 229-237.
examined the specimen. Dixon, H. N 1922 .Some new genera of mosses. J .Bot.
60:101-110.

We thank Mr, M, Thawatchai (RoyalForestDepartmenO Dixon, H. N. 1932 .0n the moss fior aof Siam, J, Siani.
awnhdo Dhre.l peK da nuys a i n S tahnet fainealed hsourtvee y s.(Ch Wie a na lgsMoai t h aUnnkiv Derrss.i tUy.i D ix oSno, cH.. ,N Na.t .1 H9i3s5t .. FSurutphpelr . 9ee: nt1ri-b5ut1io.ns to the rnoss fiora
MBi. zCu,s hTiamna an dfb rA ,eo mTpabruinwg for tohurei sr pseucgigmeesntsions wi tohn Etnhte oiddone,n- F el soefn sStieaimn. , JJ. . 1S9i8a5m. CSooncf.i, dNeantc.e H i1s"tn. iStu opsnpL ph 1y0l:o gle-n3i0e.s: an
tity of the plants a,nd Drs, A. Newton (BM) ,W. R. Buck appreach using the bootstrap. Evolution 39: 783-791.
(NY),X.He-paygr6n(H),A.Ibuw CL),C.Rausch (PC), Gangulce, H,C. 1977. Mosses of eastern India and adja-

NNIII-IE-leEcltreoncitcronic Library Service
TThheJea paJnaespeSaoncieestye Society ffoorr PPllanatnt SSyystsetmaetmicastics
September2009 AKIyAMA & TSUBOTA-Synu)lviodon ieiocarpus ,sp. nov. 95
cent regions. Fascicle 6. Privately published C,alcut- mite, Amitermes iaurensi s(Isopt eTrearm,itidae),
ta.Goffinet, with special reference to thc variety ofmound shapes.
B., W. R. Buck & A. j. Shaw. 2008. Morphology Mol. Phyl .Evo] ,42: 236-247,
and classificatiun of the Bryophyta. Ih: Goffinet, B. & RenaLLId F,. & J, Cardot ,1896[1895 ]M.usci exotici novi
A. J. Shaw (eds .B)r,yophyte Bioiogy (2n dedition), vel mixus cogniti VII, Bull, Sec. Bot. Belgique 34:
pp. 55-138 .Cambridge Univ. Press ,Cambridge. 57-78.
Guindon, S. & O. Gascuel .2003, A simple, fast ,and ac- Rohrer ,J. H. 1985. A generi crevision ofthe Hylecomia-
curate algorithm to estimate large phylogenie sby eeae. J. Hattori BoL Lab. 59: 24l-27g.
maximum likelihood .Syst .Biol. 52: 696-704. Ronquist, F. & j. P. Huelsenbeck. 2003, MRBAYES 3:
Hara, M. 2003. Vegetatio nof Doi Inthanon ,northern Bayesian phylogenetic inference under mixed mod-
Thailand .Trop. Ecol .Letters no. 51: 1･-6 .(i nJapa- els. Bioinformati c1s9 :1572-1574,
nese) Schwartz, G. 1978. Estimating the dirnension ofa model,
He, Si. [Intern reesteurce]. Checklis tof the mosses of Ann. Stat. 6: 461-464.
Thailand,http:llwww,mobot.org/MOBOT!mossf Shimodaira, H. 2002. An approximately unbiased test of
Thailand (Septem b1e3,r 2008). phylogenetic tree selection. Syst .Biol .51: 492-508.
He, Si & J. Snider. 1992. A prelirninar ysurvey ef the Shimodaira,H.2004.Approximatelyunbiasedtestsofre-
inoss genus llymphyoclon (Symphyodontace aMues,- gions using multistep-multiscale bootstrap resam-
ci). Bryobrothera 1: 283-287. pling. Ann. Stat. 32: 2616-2641.
He, Si & J. Snider .2000. A taxonomic revision of Syni- Shimodaira, H. & M. Hasegawa, 2001, CONSEL: fbr as-
pbyodon (Musc {S:ymphyodontaceae) .Bryologist sessing the confidence of phylogenetic tree selection.
103: 52-8I. Bioinformati c1s7 :1246-1247.
Horikawa, Y. & H. Ando. 1964, Contributio ntso the moss Sikes, D. S. & P. O. Lewis. 2001, PAUPRat: PAUP' im-
flora ofThailand, Nat. Life Southeas tAsia 3: 1-44, plementation ot'the parsimony ratchet, beta software,
Huelsenbeck ,J. P. & F. Ronquist. 2001. MRBAYES: version 1 ,Distribut ebdy the authors. Departnnent of
Bayesian inferenc eofphylogeny. Bioinformatics 17: Ecology and l'Ltvelutio nBairoylogy, University of
754-755. Connecticut, Storrs, USA.
Ignatov, M. S. ,T. Koponen & D. H. Norris .1999 .Bryo- Swoffbrd, D. L. 2003, PAUP": phylogellet aincalysis us-
phyte flor aef thc Huen Peninsula L.XII. Brachythe- ing parsimony ("an dother methods), version 4.
ciaceae (Musci e)xc,juding Iloinalothecium and Pa- Sinauer Associates, Sunderland.
lamocladiurn. Acta Bet. Fennica 16S: 23-72. Tamura, K,, J. Dudley, M. Nei & S. Kumar. 2007.
Kishino, H., T. Miyata & M, Hasegawa. 1990. Maximurn MEGA4: moleeular cyolutienary genetics analysis
likelihood inference ofprotein phylogeny and the ori- (MEGA) software version 4.0 .Mol, Biol .Evol ,24:
gin ofchtoroplasts. J. Mel. Eyol, 30: 151-160. 1596-1599.
Kitagawa, N. 1967a .Studie son the Hepaticae ofThailand Tanabe, A. S. 2007, KAKUSAN: a computer program to
I, The genus Bazzania, with general introductio Jn,. automatc thc sclection of a nucleotide substkution
Hattori BoL Lab. 3e: 249-270. model and the configuration of a mixed model on
Kitagawa, N. 1967b .Researche sof hepatic sin Thailand. multiiocus data. Mol. Ecol. Notes 7: 962-964.
Nat, Life SE Asia, 4: 180-183 .(i Jnapanese) Tsubota, H., E. De Luna, D. Gonzalez ,S, M, Ignatov & H,
NoguchL A. I972a. Mosses of Thailand, Lindbergia 1: Deguehi. 2004. Molecu]ar phylogenetic and ordinal
169-183. relationships based on analyses of a large-scal edata
Noguchi, A. 1972b. A revision of the gcnus ILlacrotham- set of 600 rbcL sequences of mosses. Hikobia 14:
nit{m, Kumamote J .ScL Biol. Il: 1-12. 149-169.
Noguchi, A. I973. Contributio ntso the bryo]ogy ofThai- Tsubota, H., K. Takahashi, M. Nakahara, H. Mohamed &
land. J. Hattori Bot. Lab, 37: 235-250, H. Deguchi. 2005. A simple precedure for DNA isola-
Ogawa, H,, K, Ybda & T. Klra, (196i )A, preliminary suF tionusingsmallquantitiesof1ichent]iallus.Lichenol-
vey on the vegetation of Thailand. Nat. Life SE Asia ogy 4: 2S-28.
1 :21-157. Touw, A. 196g. Miscellaneous notcs on Thai mosses. Nat.
Ozeki ,M., Y. Isagi H,, Tsubota, P. Jacklyn & D. M. j. S. HisL BulL Siam Soc. 22: 217-244,
Bowman. 2007. Phylogeograph yofan Australia tner-
ReceivedApri l21 ,2009; accaptedJttne 10 ,200P
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AppE'NDix. Taxon sampling and DDBJIEMBLfGenBank acccssion rn]mbcrs used in the present analysis (Fi g2.). Newly ob-
tainedsequenees(inbold)areshownwiththeirvoucherinformation(inorderofscientificnamc.accessionnumber,loeal-
ity ,and collector numbeD. All voucher specimens ofnewly sequcnced samples are deposite idn HYO.
Acroporium stramineum (Reinw ,& Hernsch. )M. um nemorale (Mitt ,A,) Jaeger ,AB029387; R eu,>tply1-
Fleisch. ,ABOS1225; Bracbythecium salebrosum (Hofim. rum (Card &ot Ther. )Z. Iwats .A,B024628; Plagiomnium
ex F. Weber & D. MohD Schimp., AFIS8176; Brotherella .i`u,onicu (mLindb .T.)J. Kop., AB050992; Platylt>tpnidi-
henonii (Duby )M. Fleiseh. ,AB029167; Campylophyltum um riparioides (Hedw, )Dixon, AB029385; Pleurozium
halier (iSw .ex Hedw.) M. Fleisch .A,B332270; Cratoneu- sch reberi (Will dex. Brid. M)itt. A,B024664; Ryeudo taxi-
ron.filicinu,n (Hedw .Sp)ruce ,AB095270; Ctenidiumpin- pbyUum pohliaecarpum (Sul l&. Lesq.) Z. Iwats.,
natza,n (Brot h&. Paris) Broth., AB332252; Eetropotheci- AB332271;eylaisiadeiphatristoviridis(Broth,)Afonina,
um ptychoj2]lium N. Nishim., AB332255; Entodon Tsubota & Ignatova, AB050991 [submit iase Idlypnum
challengeri (Par iCsa)rdot ,AB05e993; E. coneinnus (De tristoviride(Broth,)Paris];RAupnchostegiunzpaliidijbtium
Not.) Paris, AB332277; E. rubicrundus (Mit tA,, )Jaeger, (Mitt A.. )Jaeger A,B024944; Sanioni auneinata (Hedw.)
AB029386; Entodontopsisleucosteg(aBridW.,)R.Buek Loeske,AB332276; Setnatophyllutn (MUII,
subhumite
& Ireland, AB024635; Rmtinalis antipyretica Hedw., Hal,) M. Fleisch, subsp. .ioponieum (Broth .S)eki,
AB050949; GoUaniaJ'aponica (Cardo DAndo & Higuchi, AB039675; SF,,mplty easdpeorn (Mitt .)A. Jaeger,
AB332248; lieieropdylli ucam77ne (Hook, )M. Fleisch., AB491807, Thailand, Doi lnthanon, around the summit,
AB051218; fibmomallium japonicoaduat u,(nB,roth,)2550 m a]t., Hl Akiyama 21543 sSlvmpltyodo nerinaceus
Broth.,AB332243;Ilytocomiastrumavrenaicum(Spruce) (MittA..)Jaeger,AB491808, Thailand,Doi Inthanon,

M, Fleisch. ,AB024660; llb,locomium ,ny,gende n(sHedw.) Kew Mae Pan, 2200m alt., H. Akiyama 21562; S))implty-
Schimp. ,AB024662; Ilypnum cupressij?)"me Hedw,, oclbn leiocarpu sH. Akiyama & Tsubota, AB491804
AB032077; Ilypnum plumaq12)rme Wilson, AB029384; [submit atse tdh,mphyed ospn.], Thailand, Dei Inthanon,
LoeskeobTyum cavijlolium (Sand Leac.) M. Fleisch. ex around the summit, H/ ]klyama T7z-12 7S;l,,mpdy,odon
Broth., AB024658; Leptok4,meniufn ten"e (Hook.) scaber (Tixie St. lHe & Snider, AB49!805, Myanmar, Mt.
Schwtigr .A,B491801, Myaninar, Mt. Victoria, 2750 m Victoria, 2250 m alt., X Mhrrata et ai, 23237; Sl,impIij,odon
alt., J. Miirat aet al. 22298; Leptectadiellapsi.tur a(Mirt.)scaber, AB491S06, Thailand, Doi lnthanon, Kew Mae
M. FIeisch ,AB,491802, Myanmar, Mt. Victori a3,000 m Pan, 2200 m alt., H/ Aklyama 2J559 ;IV)uicliumpristoca-
alt., J Mtirata et al. 22380; Maerothamni"m macrocar- ly x(Mti 1H1al.. )A. Jaeger, AB071416; 7bmenmpnum ni-
pum <Reinw .& Hornsch,) M. FIeisch. ,AB491803, Thai- tens (Hedw. L)oeske, AB024676; 1}"ismegisti aiancijblia
land ,Doi Inthanon, around thc summit, 25SO m alt., Hl (Harv .B)roth,, AB051227 [submitt easd T korthaisii
Aklyama 7Zz-65 ;Adyuroclado maximowiczii (G.G .BoF (Dozy & Molk.) Broth.].
shch.) Steere & W.B. Schofiel dA.B029389; Pkrgiotheci-
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