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iSSN1346-7565 Acta Phytotax, Geobot. 62 (1): 51-59 (201 1)

Contributions to the Flora ofBorneo, 1, A New Species,

Dimorphocladon echinocarpum (, Musci),

with Unique Multicellular Exothecial Spines

HIROYUKI AKIYAMA

Miisettm qfNbture andHuman Activities, la,oigaoka-a Sanda, H),ogo Prq12]cture 669-1546, Japan. aktyama@hitohaku,1'p

Dimorphocladbn echinocaipum H, Akiyama, with conspicuous multicellular spincs on the exothecium, is described as new based on two s'pecimens frem northern East Kalimantan, Indonesia.

Key words: Bernco, bryophytes, Dimorphocladbn, East Kaljmantan, Kayan Mentarang Natienal Park,

new species, Symphyodentaceae

Northern Borneo is part of three countries; kan. Long Bawan js one ofthe main villages in- Malaysia (Sabah & SarawalO and Brunei in the side the Kayan Mentarang National Park and northwest and across the backbene range, and In- deeply involved in the WWF Heart of Borneo donesia (East Kalimantan) in the southeast. Dur- Initiative (Persoon & Osseweijer 200g), ljoined ing the 2eLh century, bryological surveys were the trips as a bryophyte specialist and coltected

conducted mostly in the Malaysian and Brunei more than 500 bryophyte specimens. In the parts of Borneo, while East Kalimantan was left course of identification, an interesting 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 diversity is very lew eamp site beside a small river at the fbot ofBatu

because of low humidity, On the other hand, a Harum, a five-day hike through the fbrests from large number 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, linear laminal cells tions are much higher, especially near the border with distinct prorulae at the upper ends, weakly with Sabah and Sarawak. differentiated alar cells, a short, double costa, and A joint botanical survey was conducted in sharply spinose capsules on smooth setae. In ad- l990 by Herbarium Bogoriense and the Univer- dition, a habitat preference for shrub leaves at sity of TIbkyo to document plant diversity in the rather sunny streamsides is also characteristic. region, especially from Long Bawan to thc peak These features suggest its close relationships to of Gunung (Mt.) Batu Harun (2169 m alt.). Long the Symphyodontaceae, especially Dineoi:phocla- Bawan is a small village at an elevation of 1000 don Dixon or 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 Syimpbyodbn Mont., 1and forests and river systems, or aone hour fiight it is described here as a new species ofDimor-

by a small plane, from the seaside city of [fara- phocladbn as discussed below in detail,

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Indonesiu, Eust Kalirnantan, Kabupaten Dimorphocladon echinocarpum H, Akiyama, Z)v7tts. Bu- lungan, Kecamatan Long Bawan, north of Long Bawan sp.nev. Figs. 1-11, 18-20, vi11agc: ln 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 1is ovoideis ad brevitcr cylindracejs, setis juxta capsulas Aklyama IC-24208 (holo- HYO; iso- BO, L, NY), argute flexis, et spinis in paginis capsularum mu]ticellu- laribusdiversum.

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FiG. 1-1O. Dimoi7)hocladon echinocai?um (all from holotype). 1. on Ieafof1iving shrub. Z. ventral side ofprostrate stem 3.deoper- showing dislalike rhizoid tu fts (arrows indLcate gemm{fereus branches with filamentous gemmae in leaf axiLs), 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 keeled segments. 10. exothecial cclls with multicellular prqcctions.

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June 2011 AKLyAMMA New Species. Dimorphocladon eckinocarpum 53

Plants green to olive-green in life, turning just below neck of capsule. Copsules upright to pale brownish yellow after drying, forming low, horizontal, ovoid to short cylindrical, deep yel- thin tufts on leaves of living shrubs. Rhizoids re- low, with short neck, 1-1,8 mm long, with mnlti- stricted to ventral surfaces of prostrate stems, cellular spines O,1-O.2 rnm long throughout; exo- tufted, smooth, brown. Stems pale green, becom- thecial cells quadrate to short rectangular, evenly ing brownish yel]ow in age, prostrate, tjghtly 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; paraphyllia and pseudopara- long; peristome double, brownish yellow, closed phyllia absent; cortical cells lax, thin walled; cpi- during dry conditions, exostome teeth 16, lanceo- dermal cells thick walled, substereid, without late belew, dorsally with narrQw medi' an furrows,

centrul strand. Branches dimorphic. Plegetativesuddenly narrowed at shoulder into filamentous

mostly simple, usually cm upper branches lessthan 1 portion,to 450 ,um inheight,basallydense- long, more or less complanately fbliate, Genemi- ly striate, margins more or ]ess appendiculate, .ferous branches similar to vegetative ones, but apically finely papiltose; endostome to 450 ptm in tapering apically, with a number of smooth height, with high basal membrane, segments lan- walled, filamentous gemmae in leaf axils; gern- ceolate, keeled, basally smooth, apically finely mae 2-2.5 mm long, Stem leaves slightly smaller pap{11ose; cilia rudimentary, 1 or absent. spores than braneh leaves. Branch leaves narrowly lan- etlipsoid, thin walled, longer axis 35-50 um long, ceolate, 1.4-1.8 mm long, more or less shallowly shorter axis 25-35 um long. concave, not decurrent at base, margins minutely Aclditional specimen examined. INDONESIA. Bor- revolute, apex long acuminate, twisted, with nco Island. East Kalimantan: Kabupaten Bulungan, Ke- short but distinct geminate 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'ama l(-24243 (HYO). laminal cells linear, ca. 100 stm long, becoming

shorter apically, prorate on upper ends on dorsal .Habitat. On leaves of short shrubs at sides of

and lower surface, walls slightly thick ; median streams in moi st secondary and primary montane laminal cells linear, walls slightly thick 80-120 , forests; quite similar to habitat of DimoFlphocla- smooth; marginal cells similar "m !ong, laminal don borneense and species of Chaetomitriune.

to inner ones but shorter; alar cells slightly difller- Disiribution, Known only from the speci- entiated, or hexagonal, 3-5 in num- quadrate ca. mens cited above; probably endemic to northern ber, pale green, not porQse. Axil]ary hairs hya- Borneo, line, un iseriate, 2-celled; basal cell quadrate,pale Distinguishing foatures, (l) Epiphyllous on apical cell elongate, smooth. brown; hyaline, short shrubs at sides of streams in montane for- Autoicous. Perigonia on stems or at prostrate ests; (2) dimorphic branches with more or less base of lateral branches, with ca. 10 antheridia complanate foliation; (3) filamentous gemmae in and few Perigonial leaves ovoid, paraphyses. leaf axils of caudate, gemmiferous branches; (4) margi.ns entire or finely serrulate apex distally, linear, slightly thick walled laminal cells; (5) low often laciniate, apiculate; costa double, 1f5 ofleaf but distinct geminate teeth at leaf margins; (6) length. Perichaetia on stems or at base prostrate short, double costae; (7) weak difTerentiation of of lateral branches, with 3-10 archegonia, with- alar cells on branch leaves and with few quadrate out Post-fertilization paraphyses. perichaetial to hexagonal cells; (8) smooth setae often sharply leaves lineaFtriangu]ar, 1.g-2.5 mm long, slight- curvedjust below capsule; (9) capsules with mul- concave, often laciniate revo- ly distally,margins ticellular spines and opercula with long, slender

Iutc and sharply serrulate, apex filamentous; cos- beak; (10) transversely striate ornamentation at ta weakly developed. Ca(zptrae cucullate, dense- base of exostome teeth; and (11) large, el]ipsoid ly setose. Setae reddish brown, ca, mm long, 8 spores to 50 ptrn long in longer axis. srnooth, straight, but sharply bent alld echinate

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Fic}. 11. Dimorphociadon eehinocarpum (a-o: all from holotype), a & b: leaves ofnormal branehcs. c: a lcaf ofgemmiferous branch. d & e: perigonial ]eaves, fi a post-fertilizaiion pcrichactial lea£ h & i/ filamentoug gemmae in leaf axils. j: apcx of branch leaf. k, apicalmaTgin ofbranch lea'E 1: mcdian laminal cells ofbranch leaE m& n: alars ofbranch leaves. o, Transverse seetien ofbranch.

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June 2011 AKIyAMA-ANewSpecies,DimorphoctadonechinocarT]um 5S

Comparison M,ith Dirnorphocladon borneense lowing morphological features with D. echino- (Figs, 12-17, 21-22) carpum; (1) dimorphic branches, (2) filamentous, Dimop:phoclaclon was established as a mono- smooth gemmae in leaf axils of gemmiferous typic genus by Dixon C1922) based on a single branches, (3) geminate teeth to base of leaf mar- specimen collected in Sarawak in north central gins, g) linear, somewhat thick walled laminal Borneo, The sole member, D. borneense Dixon, cells with proration at upper end, (5) branch was then reported from the Malay Peninsula leaves with slightly differentiated, rectangular to (Malaysia, Thailand; Dr. B. C. Tan, personal quadrate alar cel]s, (6) double costa of 1f3-1/4 of communication), the Philippines

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FTG.12-17.Dimvrphocladon borneense (tgki,yamaBorneo-32441). 12. plants on branch ef shrub (arrews indicate gemmiferous branchcs with fiLamentous gemmae in leaf axils>, 13. capsule with long slender operculum, 14. deoperculate eapsulc. 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.g., C. elegans Geh., C. torqtiescens sules with low, indistinct warts on neck. The low Bosch & Sande Lac.), but others are not (e.g,, C and indistinct warts (Figs. 21-22) are composed perlaeve Dixon) in Chaetomin'iztrn and thus can-

of several cells and thus might be rudimentary not be a separatjng feature,

stage of the well developed spines ofD. echino- It is also noteworthy that a few species of carpum. Chaetomitrium fi'om New Guinea and nearby is- Dixon (1922) regarded the geminate teeth on lands pessess echinate capsules (e,g., C. acantho- the lcaf margins o'f Dimorphocladon horneense carpum Bosch & Sande Lac.; Bartram 1961, Aki-

to be an important feature for treating the genus yama 1997). The spines on the capsules, however, as distinct from Chaetomitrium. Although spe- are merely projections, each spine arising from a cies of Synrphyodon do not have geminate tecth single exothecial ccll (Figs. 23-24), (He & Snider 2000), a number of spccies of Chae- Among the most striking featurcs separating tomitrium do have them (Akiyama 1997, Akiya- Chaetomitrium from DimoT:phocladbn are the ma & Suleiman 2001). Geminate teeth by thcm- papillose, spinose, or sometimes warty setae selves are not sufficient to distinguish Dimorpho- (Flcischer 1908, Akiyama & Suleiman 2001). On claclon from related genera. this point, it should be noted that a single species, ChaetomitFiopsis glaucocarpa ex opecimens examined Pimorphoelatlbn borneens. (Reinw. INDONESIA, Borneo, West Kalimantan: foot of Gu- Schwagr.) M. Fleisch., is separated from Chaeto- nung Ntut, 500 m alt., rL Aktyama K-31440 (HYO); Putus- mitrium by the long, smooth setae (Akiyama sibaLL, upper stream ef Sungai Nuo, 160 m alt., H/ Akiya- 2006, Fleischer 1923, Mohamed & Robinson ma KL32095 (HYO); ibid., upper reachcs ofKapuas river. 199]). According to a recent molccular phyloge- near Mata Lunai, 160 m alt. H: Aklycrma KL32477 (HYO). netic analysis 2010), Chaetomitriopsis South Kalimantan: foot ofGunung Batu Besar, 400 m (Ho glau- cocatlpa is an in-group member of Chaetomitri- alt. H. Aki.vama K-24636 (HYO). um "Tith high bootstrap probability, The smooth Comparison to ,!pecies Chaetomitrium with ojr' setae, also tbund in Dimorphocladbn, should echinate ccrpsules and Chaetomitriupsis glauco- thercfore be trcated as mere variation within carpa 23-24) (Figs. Chaetomitrium. As a result, thc relationships be- More than 110 species of Chaetomitrium have tween Dimop:phocladon, Chaetomitrium and been described, mostly from Southcast Asia. Chaetomitriopsis M, Fleisch, remain ambiguous Among them, no less than 16 specics have been and are in need of fUrther ana]ysis, reported from Borneo (Akiyama & Suleiman examined, Specimen,y ( 7iaetotnitriopsis.ol"ucocar- 2001, Su]eiman et crl. 2006). It is noteworthy that pa. INDONESIA. Borneo. East Kalimantan: Long the 16 species are strictly confined to the branch- Bawan, Bulu Kinuab,, 1070 m alt., ". Ak4yama KL24iP5 es and leaves of 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. CenLral Seram[ Tehoru, from Hunisi to Wae Heka -North Heka, 350 m ait., H/ Aklyama C-15958 dition, Chaetomitrium and DimoF]phoeladon (HYO). Seram: near Elemata-Makualaina, 10e-350 m alt., U share the fo11owing features; (1) filamentous gem- Aktyama C-9313 & C-93Z2 (both HYO). mae in leaf axils, (2) proration at upper ends of Comparison to species Symphyodon laminal cells, (3) short, double costae (usually of (Figs, 25-26) less than 113 ofleaflength in Chaetomitrium), (4) The is distributed in tem- leafmargins with geminate teeth, (5) weak differ- genus th,iuph),odon to tropical zones worldwide, Tvve species, entiation of alar regions ef branch leaves, (6) perate S. Mont. and S. copegandii Broth., short, straight hairs on calyptra, (7) long, slender perrottetii have been reportcd from Borneo, Aecording to beak of opercula, (8) striate ornamentation on He & Snider the species of exostome teeth, and (9) larger speres (Mohamed (2000), S}JmphJ,odon resemble Dimorphoctadon echinocarpum in the & Robinson 1991). As for capsule posturc, some

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57 June 2011 AICIYAMA ANew SpeCies，Dimorphoc 〜α ゴ〜）n ecinnoca ηpum

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FIG 18−26 Splnes o 墅1 capsules Dmvorphocladon echtnocar πwTn 18−20 h 〔｝k〕type O borneense 21−22 Akiyama 1（3247 ア （ ， ）， （ ， ，

− West Kal皿 antan ）， Chaetemttrium acanthocarl フ跚 （23 24， Akiyama C 159S8， Serdm Is1乱nd ）、md S）mphyodon gca ろer

T 正xlel SHc ＆ Snider 25−26 IJ Akl crma et a ／ 〃 7 Thalland Dol Inthanon 18 ＆ 19 closeup ofneck of capsule and （ ） 〔 ， ｝ ， ， ） ． しurvcd ， roughly し chinate upperrnost portion ofa ＄eta 20 twomul 亡lcel ］ular spmes w ith clongate ceLls on capsule walls 21 ＆ 22ne し k portlon of し apsuLes ， note lew， muhl し ellular warLg 23 ＆ Z5 deopelculdtc capguleb 24 ＆ 26 un1 しcllular spines on capsuL し walls

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(1) proration of 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 ferences also suggest a rather distant relation- leaves with distinctive serrations 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 discussed above, thc relationship of Di- Sympbyodbn diffe･rs from Dimorphocladon in fnoiphoctadon echinocarpune with other mem- sporophytic features and habitat preference as bers of the Symphyodontaceae cannot be deter- fbllows; (1) exostome teeth lifiear-lanceolate, out- 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 its systematic posj- and linear segments, (3) smaller spores ranging tion, especially in regard to Chaetomitrium. 13･-20 ptm in diameter, and g) preferring some- Dimoivhoetaclon echinocaT:pum and related what shaded fbrest floors and edges of fbrests (He species and genera all share the short, double ces-

& Snider 2000). In addition, each spine on the tae, lipear and more or less thick walled lamina! capsule walls in syneph.vodon is a mere projection cells with proration at the upper end, slight differ- firorn a single exothecia] cell (Figs. 25-26), as in entiation of alar cells, and double peristomes.

Chaetomit"ium acanthocaizpum, and differs con- They may be distinguished by the foTlowing key.

siderably from the multicellular spines ofDimor-

Keys to Di.mor;phoeladon echinoearpum and relatcd taxa

1a. Setae spinose, setose, or highly mammillose .,.,...... v.."...,.,"..,,...... ,....,..-..,,. Chaetomitrium spp lb. Setae smooth (often mamrnillose er papillose apically)..,..."..,,...".,,,.,.k,,,...... ,."...,,,.."...,..,,..".,,,..,...... 2 2a,Capsulesspinoseorwarty...... ,..-...... ,..,,...,...,,,,...... ,..."..,,,..v.,,,,,,.....,,,,...H...,..",,,,....,...... ,,,.H.,,,.,...... ,.,3 2b.Capsulessmooth-...,...... "...,,.H...,..."..,...... ,...-..,..,-"."",.,...... -,..,,.H4 3a, Exostome tceth papil]ose at base; endostome with 1inear segments and low basal membrane; spincs on cap- sules unicellular, a mere cuticle prejection of cell wall .....,.,,...,,...... ,,...... ,...... ,...... ,.,... Symp1tyodon spp. 3b. Exostome teeth densely striate at base; endostome with lanceolatc segmcnts and high basal membrane; spineoncapsu]esmulticellular"..,,,.".,...,.H-..,,,..".,.,,,."...,,.."..,.,..-..,...,...,.-.DimorphocladonechinocaFT)um 4a, Plants complanately fbliate; fiLamcntous gemmae present in leaf axils of gemmiforous branches; teeth of Leaf margins gcminate; setae to 20 mm long...,,.-....,,,..m...... ,....-...... "..,,,.. Dirnorlphocladon bomeense 4b. Plants roundly fbliate, squarrose. Gemmac absent; tc-cth of leaf margins simple; setae to 40 mm long .....-..,,,,.,.,"...... -,.,,...,...... ,,...... ,...... ,.,...... ,,,...... ,...... ,.,,,,...... ,,,.-,.,,..H...ChaetotnitFiopsi,yglaucocarpa

I thank Dr. M. Kato (Nationa] Museum ofNature and Sci- by Indonesian-Japancse Botanical Expeditions VIII, ence, Tokyo) whe offered me the chance to join the eo- Meteoriaceae, Hookeriaceac, and Trachypodaceae, operative botanical expeditions to eastern, western, and Nature and Human Activities 2i 9-3 1.

southern Kalimantan held by Univcrsity of Tokye and Akiyama, H. 2006. Ncw synonymy of Chaetomitriopsis Herbarium Bogeriense (lndonesian Institute of Scienees gtauc'occrrpa (Hypnaccae, Muscj), Trop. BryeL, 2g: LIPI), Dr, W, R. Buck Yk)rk Botanical Garden) fbr 61.Akiyama, (New 'Taxonomical linguistic corrections, and Dr. H. Nagamasu (Kyote Uni- H. 2009. studies of. ofSc- versity Museum) far his kind help in preparing ihe Latin ram and Ambon (Moluecas, East Malesiaj colleeted diagnosis. by Indonesian-Japanese Botanical Expeditions, X. Bartramjaeeao, Brachytheciaceae, Bryaceae, Dit- References richaceae, Erpodiaceae, Hylocomiaeeae, Hypnaceac ({nparti,Mniaceac,Orthotriehaccae,Regmatodonta- ccae, and Splachnaceae with correctionsXaelditions to Akiyama, H, 1997, Taxonemic studies of mosses of Se- the previous repons. Humans and Nature 20: 15-28. rain and Ambon East Malesial collected (Moluccas, Akiyama, H, & M. Sulciman. 2001 . Taxonomical notcs on

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June 2011 AKiy,NMA-A New Species, Dimorphoekidon echinocai7pum S9

lhc genus C7taetomitrittm (Hookeriaceae, Musci) of Mohamed, I'I. & H. Robinson. 1991. Ataxonomic revision Borneo. Hikobia 13: 491-509. of the mess family Hookeriaceae and Hypopterygia- Bartram, E. B. 1961. Low aLtitude mosses from northeast- ceae in Malaya. Smithsonian Contr. Bot, 80: 1-44 t ern New Guinca. Brittonia 13: 368-380. i---iv.

Dixon, T-I. N. 1922, Some new genera of mosses. j. Bot. Persoon, G. A. & M. Osseweijer. 2008. Reflections on the 60: 101-110. Heart ef Borneo. Tropenbos InteTnational, Wagenin- Fleischer, M. I908. Die Musci der Flora von Buitenzorg, gen. (http://www,tropenbos.orgftbi-publicationsl voL 3. E. J. Brill, Leiden. decuments!TBIntSeries-24.pdf). Fleischer, M, 1923. Die Musci der Flora von Buitcnzorg, Schultz-Motel, W. 1963. Vt)rltiufiges Verzeichnis der

voi. 4. E. J. Brill, Leiden. Laubmoose von Ncuguinca. Willdenowia 3: 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 firom Born ¢ Q. J, Hattori Ho, B.-C. 2010. Evelution and Diversification of the Botanical Lab. 99: 107-183.

Hookeriales () with Emphasis on Disticho- Tan, B. C. 1993. Noteworthy range extension ofMalesian plyUum (Daitoniaceae) and Its Allied Genera. Dis- mosses. J. Hattori Bot. Lab. 74: 227=233. sertation,Friedrich-Wilhelms-Universittit,Bonn.

ReeeivedApril 5, 20".' accepted Miry 11, 20.ll

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