Contributions to the Moss Flora of Borneo, 1. A New Species, Dimorphocladon echinocarpum (Symphyodontaceae, Musci), with Unique Multicellular Exothecial Spines

2011·3.4 MB·English

by  Akiyama, Hiroyuki

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iSSN1346-7565 Acta Phytotax, Geobot. 62 (1 )5:1-59 (20 11)
ContributionstotheMoss Flora 1,A New Species,
ofBorneo,
Dimorphocladon (SymphyodontaceMuasec,i),
echinocarpum
with Unique Multicellula Erxothecial Spines
HIROYUKIAKIYAMA

Miisett mqfNbture andHuman Activitie sl,a,oigaoka -Saanda, H),o gPorq12]ctu 6r6e9-1546 ,Japan.
aktyama@hitohaku,1'p
Dimorphocladbn echinocaipum H, Akiyama, with conspicuous multicellular spincs on the exothecium,
is describe das new based on two s'pecimens frem northern East Kalimantan ,Indonesia.
Key words: Bernco, bryophytes, Dimorphocladbn, East Kaljmantan, Kayan Mentarang Natiena lPark,
new species, Symphyodentaceae
Northern Borneo is part of three countries; kan. Long Bawan js one ofthe main villages in-
Malaysia (Saba &h SarawalO and Brunei in the side the Kayan Mentarang Nationa lPark and
northwest and across the backbene range, and In- deepl yinvolve din the WWF Heart of Borneo
donesi (aEas Ktalimantan) in the southeast. Dur- Initiativ (ePerso &o nOsseweijer 200g), ljoined
ing the 2eLh century, bryologica lsurveys were the trips as a bryophyte specialist and coltected
conducted mostly in the Malaysian and Brunei more than 500 bryophyte specimens. In the
Borneo, EastKalimantan left identification,interesting
parts of while was course of an moss was
nearly untouched, Eastern part ofEast Kaliman- found among the collections,
tan mostly consists ofa vast range oflowland be- The moss was collected at 1000 m alt. near the
low 100 m and bryophyte diversit iys very lew eamp site beside a small river at the fbot ofBatu
because of low humidity ,On the other hand, a Harum, a five-da yhike through the fbrest sfrom
larg enumber ofbryophytes are to expected in the Long Bawan. It is characterized by geminate
western part of East Kalimantan, where eleva- teeth on the leaf maTgins, linea rlaminal cells
tions are much higher ,especially near the border with distinc ptrorula eat the upper ends, weakly
with Sabah and Sarawak. differentia taleadr cells, a short, double costa, and
A join btotanica lsurvey was conducted in sharply spinose capsules on smooth setae. In ad-
l990 by Herbarium Bogoriense and the Univer- dition ,a habita tpreferenc efor shrub leave sat
sity of TIbkyo to document plant diversi tiyn the rather sunny streamsides is also characteristic.
region, especially from Long Bawan to thc peak These feature ssuggest it sclose relationships to
of Gunung (Mt. B)atu Harun (216 9m alt.). Long the Symphyodontaceae ,especially Dineoi:phocla-
Bawan is a small village at an elevation of 1000 don Dixon or Chaetomitrium Dozy & Molk. De-
m. It is situated near the border of the Malaysian spite the superficial resemblance ofthe spines on
state of Sabah, about a one week walk across 1ow- the capsule walls to those of Syimpbyod Mbonnt.,
1and forest asnd river systems, or aone hour fiight it is describe dhere as a new species ofDimor-
by a small plane, from the seaside city of [fara- phocladbn as discusse dbelow in detail,
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S2 AetaPhytotax.Geobot. Nbl,62

Dimorphocladon echinocarpum H, Akiyama, Z)v7 tIntdson.esiu, Eust Kalirnantan, Kabupaten Bu-
sp.nev. Figs .1-11, 18-20, lungan ,Kecamatan Long Bawan, north of Long Bawan
vi11ag cl:n thc vicinity of Bulu Kinuab, fbot ofBatu Har-
Affine Dimorphoctado borneensi Dixon. ,sed capsu- un, ca. 1070 m alt., 4005'N ,1150SO'E ,20 Sep. t990, HL
1i sovoideis ad brevitcr cylindracejs, setis juxt acapsulas Aklyama IC-24208 (holo H-YO; iso- BO, L, NY),
flexis, in
argute et spinis paginis capsularum mu]ticellu-
laribusdiversum.

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FiG.1-1O. Dimoi7)hocladon echinocai?um (a lflrom holotype) .1. plants on Ieafof1ivin gshrub. Z. ventral side ofprostrate stem
showing dislalik erhizoid tu fts (arro iwndsLcat egemm{fereus branches with filamentous gemmae in leaf axiLs), 3. deoper-
cul atc capsulc with closed peristomes in dry cQndition. 4. echinate capsule with a cucullatc, sctosc calypLra. 5. closcup of
young calyptra. 6. denbe]y echin ale capsules and smooth epercula with long, slender bcaks. 7. outer surface ol'perisLome.
8. transversely st-ate ornamentation on lower portion of exostoTnc teeth 9. inner surfacc of endostome with high basal
mcmbranc and keele dsegments. 10. exothecial cclls with multicellular prqcctions.
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June 2011 AKLyAMMA New Species .Dimorphocladon eckinocarpum 53
Plant sgreen to olive-green in life ,turning jus btelow neck of capsule. Copsules upright to
pale brownis hyellow after drying ,forming low, horizonta lo,void to short cylindrical, deep yel-
thin tufts on leaves of livin gshrubs. Rhizoids re- low, with short neck, 1-1, 8mm long ,with mnlti-
stricted to ventral surfaces of prostrat estems, cellular spines O,1-O.2 rnm long throughout; exo-
tufted ,smooth, brown. Stems pale green ,becom- thecial cells quadrat eto short rectangular, evenly
ing brownish yel]ow in age, prostrat etj,ghtly at- thick walled, smooth; stomata absent; annuli
tached to substrate by rhizoids, densely and pin- present ;opercula long rostrate, smooth, to 1 mm
nately branchcd ;paraphylli aand pseudopara- long ;peristome double ,brownish yellow ,closed
phylli aabsent; cortical cells lax ,thin walled; cpi- during dry conditions, exostome teeth 16, lanceo-
dermal cells thick walled, substereid, without lat ebelew, dorsall ywith narrQw medi' an furrows,
centrul strand. Branches dimorphic .Plegetativesuddenly narrowed at shoulder into filamentous
branches mostly simple, usually less than 1 cm upper portion, to 450 in height b,asall ydense-
,um
long, more or les scomplanately fbliat eG,enemi- ly striate, margins more or ]ess appendiculate,
.ferous branches similar to vegetative ones, but apically finel ypapiltos een;dostome to 450 ptm in
tapering apically, with a number of smooth height ,with high basal membrane, segments lan-
walled, filamentou sgemmae in leaf axils; gern- ceolate, keeled ,basall ysmooth, apically finely
mae 2-2.5 mm long ,Stem leave sslightly smaller pap{11os eci;lia rudimentary, 1 or absent. spores
than brane hleave sB.ranch leave snarrowly lan- etlipsoid, thin walled, longer axis 35-50 um long,
ceolate, 1.4-1. 8mm long ,more or les sshallowly shorter axis 25-35 um long.
decurrent base,
concave, not at margins minutely Acldition asplecimen examined. INDONESIA. Bor-
revolute, apex long acuminate, twisted, with nco Island. East Kalimantan: Kabupaten Bulungan, Ke-
short but distin cgteminate teeth throughout; cos- camatan Long Bawan; in the vicinity of Tadur Bangar,
ta double ,reaching 113-1" of leaf length ;upper 1050-1100 m aLt, 04015'N, 1150soiE, 20 Scp. 1990, H.
Akty'amal(-2424(3HYO).
laminal cells linea rc,a. 100 stm long ,becoming
shorter apically, prorat eon upper ends on dorsal .Habitat .On leave sof short shrubs at sides of
surface, walls slightly thick ; median and lower streams in moi st secondary and primary montane
laminal cells linear ,walls slightly thick , 80-120 forest sq;uite similar to habita tof DimoFlphocla-
"m !ong ,smooth; marginal lamina lcells similar don borneense and species of Chaetomitriune.
to inner ones but shorter; alar cells slightly difller- Disiribution ,Known only from the speci-
hexagonal, 3-5in
entiated, quadrate or ca. num- mens cited above; probabl yendemic to northern
ber, Axil]aryhairshya-
pale green, not porQse. Borneo,
line u,n iseriat 2e-,celled ;basal cell quadrate, pale Distinguishing foatur e(sl, )Epiphyllou son
brown; apical cell elongate, hyaline ,smooth. short shrubs at sides of streams in montane for-
Autoicous .Perigonia on prostrate stems or at ests; (2 )dimorphic branches with more or less
base of latera lbranches ,with ca. 10 antheridia complanate foliatio (n;3 f)ilamentou gsemmae in
and few paraphyses .Perigonial leaves ovoid, leaf axils of caudate, gemmiferous branches ;(4)
margi.ns entire or finel yserrulate distal laype,x linea rs,lightly thick walled laminal cells; (5 )low
often laciniat aepi,culate; costa double ,1f5 ofleaf but distin cgteminate teeth at leaf margins; (6)
length .Perichaetia on prostrat setems or at base short, double costae; (7 )weak difTerentiat iofon
of latera blranches ,with 3-10 archegonia, with- alar cells on branch leave sand with few quadrate
out paraphyses. Post-fertiliza tpieornichaetial to hexagonal cells; (8 )smooth setae often sharply
leave slineaFtriangu] a1r.,g-2. 5mm long ,slight- curvedjust below capsule; (9 )capsules with mul-
ly concave, often laciniat edistall mya,rgins revo- ticellula srpines and opercula with long, slender
Iut cand sharply serrulate, apex filamentou sco;s- beak; (10 )transversel ystriate ornamentation at
ta weakly developed .Ca(zptra ecucullate, dense- base of exostome teeth ;and (11 l)arge ,el]ipsoid
ly setose. Setae reddish brown, ca, 8 mm long, spores to 50 ptr nlong in longer axis.
srnooth, straight, but sharply bent alld echinate
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Fic} .11. Dimorphociadon eehinocarpum (a-o a:ll from holotype )a, & b: leaves ofnormal branehcs. c: a lcaf ofgemmiferous
branch. d & e: perigonia l]eaves ,f ia post-fertilizai picornichactia llea£ h & i /filamentou ggemmae in leaf axils. j :apcx
of branch leaf. k, apicalmaTgin ofbranch lea' E1 :mcdian laminal cells ofbranch leaE m& n: alars ofbranch leaves .o,
Transverse seetien ofbranch.
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June2011 AKIyAMA-ANewSpecies,DimorphoctadonechinocarT]um 5S

Comparison Dirnorphocladonborneense lowing features D.
M,ith morphological with echino-
(Fig1s2,-17,21-22) (1)dimorphicbranches,(2)filamentous,
carpum;
Dimop:phoclaclon was established as a mono- smooth gemmae in leaf axils of gemmiferous
typic genus by Dixon C192 2b)ased on a single branches ,(3 )geminate teeth to base of leaf mar-
specimen collected in Sarawak in north central gins ,g) linea rs,omewhat thick walled laminal
Borneo, The sole member, D. borneense Dixon, cells with proratio nat upper end, (5 )branch
was then reported from the Malay Peninsula leave wsith slightly differenti artecetadng,ular to
(Malays iTah,ailand ;Dr. B. C. Tan, personal quadrate alar cel]s, (6 )double costa of 1f3-1/ 4of
communication), the Philippines <Ta n1993), Se- leaf length, (7 )opercula with long and s]ender
ram (Akiyam a1997 ,2009), and New Guinea beak, (8 )non-collenchymatous exothecial cells
(Schultze-M lo9t6e3l) ,The species grows on the smooth and thick walled, (9 )striate ornamenta-
branches and livin gleave sof shrubs beside tion at base of exostome teeth, and (1 0h)igh basal
streams and small rivcrs in forest sA.ccording to membrane of endostome with keeled segments,
my fiel dobservations, D, borneense is usually Dimoi:phocladb nborneense diffbr sfrom D.
found in well preserve dor secondary riparian echinocarpum in the fbllowin gfeature s;(1)
thickets in lowlands and lower montane forest sin growing at lower etevations (100-8 0m0 on the
southern and western Borneo. Malay Peninsulla ,Borneo and Seram; Akiyalma
Dimorphocladbn borneense shares the fbl- 2009), (2 )longer setae (15-2 m0m long) ,(3 )erect,
W
ee
es
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FTG. 1 2-17. Dimvrphocladon borneens e(tgki,yamaBorneo- 3122.4 p4la1n)t.s on branch ef shrub (arre wisndicat egemmiferous
branchcs with fiLamentous gemmae in leaf axils>, 13. capsule with long slender operculum, 14. deopercula teaepsulc. 15.
outer surface ofexostorne. 16. inner surfaee ofendostome with highbasal membrane andkeeled segmcnts, 17. exotheeial
cells.
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cylindrical capsules, and e) smooth walls ofcap- are erect (e. gC.. ,elegans Geh., C. torqtiescens
sules with low, indistinc twarts on neck. The low Bosch & Sande Lac.) ,but others are not (e. gC,,
and indistinc twarts (Fig s21.-22) are composed perlaeve Dixon) in Chaetomin'izt arndn thus can-
of several cells and thus might be rudimentary not be a separatjng feature,
stage of the well develope sdpines ofD. echino- It is also noteworthy that a few species of
carpum. Chaetomitrium fi'om New Guinea and nearby is-
Dixon (192 2re)garded the geminate teeth on lands pesses sechinate capsules (e, gC.. ,acantho-
the lcaf margins o'f Dimorphocladon horneense carpum Bosch & Sande Lac.; Bartram 1961, Aki-
to be an important featur efor treatin gthe genus yama 1997) .The spines on the capsules, however,
as distinc ftrom Chaetomitrium .Although spe- are merely projection esa,ch spine arising from a
cies of Synrphyodon do not have geminate tecth single exothecial ccll (Fig s2.3-24),
(He & Snider 2000), a number of spccies of Chae- Among the most striking featurc sseparating
tomitrium do have them (Akiyam a1997, Akiya- Chaetomitriu mfrom DimoT:phocladb nare the
ma & Suleiman 2001) .Geminate teeth by thcm- papillos e,spinose, or sometimes warty setae
selves are not sufficient to distingui sDhimorpho- (Flcisc 1h9e0r8, Akiyama & Suleiman 2001) .On
claclon from related genera. this point ,it should be noted that a single species,
opecime nexsamined Pimorphoelatlbn borneens. ChaetomitFiopsis glaucocarpa (Rein w.ex
INDONESIA, Borneo, West Kalimantan: foot of Gu- Schwagr. )M. Fleisch .i,s separated from Chaeto-
nung Ntut ,500 m alt., rL Aktyama K-31440 (HYO) P;utus- mitrium by the long, smooth setae (Akiyama
sibaLL, upper stream ef Sungai Nuo, 160 m alt., H/ Akiya- 2006, Fleischer 1923, Mohamed & Robinson
mnSeaoaru Kt MLha 3tK2aa0l 9iL5 um(naaHnitY, aO1n6): 0 i; fmboi odatl . tou.,fp GHpue:nr uAn kgrleaycchr cmBsaa tKoLufK 3aB2pue4a7ss7ar (,4 Hr0iY0veO rm.). n1e9ti9c] )a n.Aaclycsoisrdi (ngH ot o2 0a1 r0e)c ,eCnhta meotloccmuiltarri opphsy iglsloague--
alt. H. Aki.vama K-24636 (HYO). cocatlpa is an in-grou pmember of Chaetomitri-
um "Tith high bootstra pprobabili tTyh,e smooth
Comparison to ,!pecie soj rCh'aetomitrium with tbund in Dimorphocladbn,
setae, also should
echinate ccrpsules and Chaetomitriups igslauco- thercfore be trcated as mere variation within
(Fig2s3.-24)
carpa Chaetomitrium .As a result, thc relationships be-
More than 110 species of Chaetomitrium have tween Dimop:phocladon ,Chaetomitriu m and
been described ,mostly from Southcast Asia. Chaetomitriops iMs, Fleisch r,emain ambiguous
Among them, no less than 16 specics have been and are in need of fUrther ana]ysis,
reported from Borneo (Akiyam a& Suleiman
2001, Su]eiman et crl. 2006). It is noteworthy that Specimen ,exyamined, ( 7iaetotnitriops i.sol"ucocar-
the 16 species are strictly confined to the branch- pBaaw. anI,ND OBNuElSuI AK.inua bB,o, r1n0e7o0. mE aasltt. ,"K.a lAikm4aynatmaan: K L2L4oinPg5
es and leave sof shrubs growing along streams in (HYO).
wcll preserved, lowland and lowcr montane fbr- C7iaetomitriuntacanthocarpuin.INDONESIA,Ma-
ests, as are the species ofDimorphocladon. In ad- luku. CenLra lSeram[ Tehoru, from Hunisi to Wae Heka
dition ,Chaetomitrium and DimoF]phoeladon Heka, 350 m ait. ,H/ Aklyama C-15958 (HYO) .-North
share the fo11owin gfeature s(;1 f)ilamentou gsem- ASkertayma:m aneCar-9 3E1l3e&matCa--9M3aZk2ua(lbaointahH,Y O)1.0e-350 m alt., U
mae in leaf axils, (2 )proratio nat upper ends of
laminal cells, (3 )short, double costae (usuallyComparison to species of Symphyodon (Figs,
less than 113 ofleaflength in Chaetomitrium) (,4) 25-26)
leafmargin swith geminate teeth ,(5 )weak differ- The genus th,iuph), oids odnistribut eidn tem-
entiation of alar regions ef branch leaves ,(6) perate to tropica lzones worldwide, Tvve species,
short, straight hair son calyptra, (7 )long ,slender S. perrotteti iMont. and S. copegandii Broth.,
beak of opercula, (8 )striate ornamentation on have been reportcd from Borneo, Aecording to
exostome teeth ,and (9 )large rsperes (Mohamed He & Snider (2000 )th,e species of S}JmphJ,odon
& Robinson 1991) .As for capsule posturc ,some resemble Dimorphoctadon echinocarpum in the
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(1 p)roratio onf upper ends of 1inear laminal cells, phocladbn echinocar;pum, As shown in a molecu-
(2 )short, double costae, (3 )weak differentiation lar analysis (Ho 2010), thcse morphological dif
of a}ar cells of branch leaves ,g) margins of ference salso suggest a rather distan trelation-
leave swith distincti vseerrations throughout, (5) ships between Dimorphoctadon and S?mph.y-
smooth setae, (6 )echinate to highly papillose Odoll.
capsule walls, and (7 )epiphytic nature growing
on branches Qf short shrubs and undergrowth, As discusse dabove, thc relationship of Di-
Sympbyodbndiffe･rfsrom Dimorphocladon in
fnoiphoctadon echinocarpune with other mem-
sporophytic feature sand habita tpreferenc eas bers of the Symphyodontaceae cannot be deter-
fbllows (;1 )exostome teeth lifiear-lanceo louatt-e, mined morphologicaily. Although tentatively
er surfaee at base sparsely papillose ,(2 )endo- classified under DimorphocXadon, molecular
stome rudimentary, with low basal rnembranes analyses are needed to settle it ssystematic posj-
and linea rsegments, (3 )smaller spores ranging tion, especially in regard to Chaetomitrium.
13･-2 0ptm in diameter ,and g) preferrin sgome- Dimoivhoetaclon echinocaT:pum and related
what shaded fbres tfloor sand edges of fbrest (sHe species and genera all share the short, doubl eces-
& Snider 2000). In addition, each spine on the tae, lipea rand more or les sthick walled lamina!
capsule walls in syneph.vo ids oa mnere projection cells with proratio ant the upper end, slight differ-
firo ran single exothecia] cell (Fig s25.-26), as in entiation of alar cells, and double peristomes.
Chaetomit"ium acanthocaizpum, and differ cson- They may be distinguish ebdy the foTlowin gkey.
siderably from the multicellular spines ofDimor-
Keys to Di.mor;phoelad oecnhinoearpum and relatcd taxa
1a .Setae spinose, setose, or highl ymammillose .,.,.......................v.."... ,C.h,a"e.to.m,i,t.r.i.u.m. .s.pp.,....,..-..,,.
lb. Setae smooth (oft meamnrnillose er papillos aepically)..,..."..,,...".,,,.,.k,,,.............,."...,,,.."...,. .2,,..".,,,..,.......
2a,Capsulesspinoseorwarty..........,..-.........,..,,...,...,,,,.......,..."..,,,..v.,,,,,,.....,,,,...H...,..",,,,....,......,,,.H.,,,.,.......,.,3
2b.Capsulessmooth-...,............................................................."...,,.H...,..."..,.............,...-..,..,-"."",.,.......-,..,,.H4
3a, Exostome tceth papil]ose at base; endostome with 1inea sregments and low basal membrane; spincs on cap-
sules unicellular, a mere cuticle prejecti oofn cell wall .....,.,,...,,......,,......, .S.y.m.p..1.t.y.o.d. .osp.np..,.......,.,...
3b. Exostome teeth densely striate at base; endostome with lanceola tscegmcnts and high basa lmembrane;
spineoncapsu]esmulticellular"..,,,.".,...,.H-..,,,..".,.,,,."...,,.."..,.,..-..,...,...,.-.DimorphocladonechinocaFT)um
4a, Plants complanately fbliat ef;iLamcntou gsemmae presen tin leaf axils of gemmiforous branches ;teeth of
Leaf margins gcminate ;setae to 20 mm long...,,.-....,,,..m.........,....- .D.i.r.n.o.r.l.p.h.o.c"l.a.d,o,n, b.o.meense
4b. Plants roundly fbliate ,squarrose. Gemmac absent; tc-ct hof leaf margins simple; setae to 40 mm long
.....-..,,,,.,.,"........-,.,,...,.........,,........,.........,.,......,,,.......,........,.,,,,........,,,.-,.,,..H...ChaetotnitFiopsi,yglaucocarpa
I thank Dr. M. Kato (Natio nMuas]eum ofNature and Sci- by Indonesian-Japancse Botanica lExpeditions VIII,
ence, Tokyo) whe offered me the chance to joi tnhe eo- Meteoriaceae, Hookeriaceac, and Trachypodaceae,
operative botanica lexpeditions to eastern, western, and Nature and Human Activitie s2i 9-3 1.
southern Kalimantan held by Univcrsity of Tokye and Akiyama, H. 2006. Ncw synonymy of Chaetomitriopsis
Herbarium Bogeriense (lndonesi Iannstitut eof Scienees gtauc'occrrp a(Hypnacc aMeus,cj) ,Trop. BryeL ,2g:
LIPI), Dr, W, R. Buck (New Yk)rk Botanical Garden) fbr 61.Akiyama, 'Taxonomical
linguist icocrrections, and Dr. H. Nagamasu (Kyot eUni- H. 2009. studies of. mosses ofSc-
versity Museum) far his kind help in preparin gihe Latin ram and Ambon (Molueca sEa,st Malesiaj colleeted
diagnosis. by Indonesian-Japanese Botanical Expedition s,X.
Bartramjaeeao ,Brachytheciaceae, Bryaceae, Dit-
References richaceae, Erpodiaceae H,ylocomiaeeae, Hypnaceac
({nparti,Mniaceac,Orthotriehaccae,Regmatodonta-
ccae, and Splachnaceae with correctionsXaelditions to
Akiyama, H, 1997 ,Taxonemic studies of mosses of Se- the previou srepons. Humans and Nature 20: 15-28.
rain and Ambon (Molucc aEsas,t Malesial collected Akiyama, H, & M. Sulciman .2001 . Taxonomical notcs on
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
June 2011 AKiy,NMA-A New Specie sD,imorphoekidon echinocai7pum S9
lhc genus C7taetomitri t(tHmookeriace aMues,ci) of Mohamed, I' I&. H. Robinson .1991 .Ataxonomic revision
Borneo. Hikobia 13 :491-509. of the mess famil yHookeriacea eand Hypopterygia-
Bartram, E. B. 1961 .Low aLtitude mosses from northeast- ceae in Malaya. Smithsonian Contr. Bot, 80: 1-44 t
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60: 101-110. Heart ef Borneo. Tropenbos InteTnational W,agenin-
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voL 3. E. J. Brill ,Leiden. decuments!TBIntSeries-24.pdf).
Fleische rM,, 1923 .Die Musci der Flora von Buitcnzorg, Schultz-Motel, W. 1963. Vt)rltiuf igVeesrzeichni sder
voi. 4. E. J. Brill ,Leiden. Laubmoose von Ncuguinca . Willdenowi a3: 399･･･
He, S. & J. A. Snider ,2000, A taxonomic revision of Sym- 549,
ph.vodon (Musci :Syrnphyodontaceae). Bryologist Suleiman ,M., H. Akiyama & B. C. Tan, 2e06. A revised
103: 52-81. catalogue ofmosses reported firo mBorn¢Q. J, Hattori
Ho, B.-C. 2010. Evelution and Diversificatio nof the Botanica lLab. 99: 107-183.
Hookeriales (Bryopsid wait)h Emphasis on Disticho- Tan, B. C. 1993. Noteworthy range extension ofMalesian
plyUum (Daitoniace aande) Its Allied Genera. Dis- mosses. J. Hattor iBot. Lab. 74: 227=233.
sertation,Friedrich-Wilhelms-Universittit,Bonn.
ReeeivedApril 5, 20".' accepted Miry 11, 20.ll
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