The Bryophytes of Sabah (North Borneo) with Special Reference to the BRYOTROP Transect of Mount Kinabalu. IV. () Author(s): Johannes Enroth Source: Willdenowia, Bd. 18, H. 2 (Feb. 22, 1989), pp. 529-554 Published by: Botanischer Garten und Botanisches Museum, Berlin-Dahlem Stable URL: http://www.jstor.org/stable/3996371 . Accessed: 03/05/2011 04:57

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http://www.jstor.org Willdenowia18 - 1989 529

JOHANNES ENROTH

The bryophytesofSabah (NorthBorneo) with specialreference to the BRYOTROP transectof Mount Kinabalu. IV.Leucobiyaceae (Bgyopsida)

Abstract Enroth,J.:The bryophytesof Sabah(North Borneo) with specialreference to the BRYOTROPtransect ofMount Kinabalu.IV. Leucobryaceae (Bryopsida). -Willdenowia 18: 529-554. 1989.-ISSN 0511-9618. In Borneothe familyLeucobiyaceae (in the old sense)is representedby the generaLeucobryum (8 species), Schistomitrium(3), Cladopodanthus(4), Leucophanes(4), Arthrocormus (1), Exostratum(2), and Octoblepha- rum(1). stenophyllum is newly recorded for Borneo. 16 namesare reduced to synonymy.Leu- cobryumarfakianum, L. papuense,and L. subsanctumare lectotypified. All species dealtwith are briefly describedand illustrationsare cited.The and ecologyof eachspecies are discussed, and their geographicalranges as well as Bomean altitudinalranges are given.

Introduction

Leucobhyaceaesensu Brotherus(1924) is nowadaysconsidered an artificialassemblage of sharingsome superficialsimilarities. It is noteworthythat Dixon &Jameson(1896) already placed Leuco- bryumHampe in the .Somewhat later Fleischer (1904) segregated the familyLeucophanaceae from the Leucobryaceae,mainly due to peristomialdifferences. Fleischer's Leucophanaceae included the generaOctoblepharum Hedw., Leucophanes Brid., Arthrocormus Dozy& Molk.,and Exodictyon Card. How- ever,this familyhas not gained generalacceptance among bryologists,and Ellis (1985) formallytrans- ferredthese genera, and his newExostratum L. T. Ellis, which he segregatedfrom Exodictyon, to the Calym- peraceaeas a separatesubfamily Leucophanoideae. In general terms, Ellis' solution seemscorrect, but I still find Octoblepharumslightly anomalous in this company;its actualplacement in the systemob- viously requiresfurther study. The remaininggenera of the classicalLeucobryaceae, namely Leucobryum (incl. CardotiaTher.), Ochro- bryumMitt., Schistomitrium Dozy&Molk., and CladopodanthusDozy& Molk., are probably best placed in the Dicranaceae.To divide the Leucobryaceaein the mannerdescribed above was alreadysuggested by Andrews(1947), and this is also the modernview (e.g. Crosby& Magill 1981). The reasonwhy these mosseswere lumped togetherin one familyis the dominanceof large,thin walled,empty hyalocysts(or leucocysts)above the chlorophyll-containingchlorocysts, which form in the leaves 1-3 layerssandwiched between the hyalocysts.This gives the mosses a distinctwhitish or bluish-greencolor, which has often been referredto as the "leucobryoidhabit". However, this kindof leaf structurehas definitelyemerged independently in differentevolutionary lineages; this is witnessedby 530 Enroth:The bryophytesof Sabah(North Borneo) IV.

the fact thatthe genus TheriotiaCard. in the Buxbaumiaceae(orDiphysciaceae, depending on the interpre- tation) has an analogousleaf structure. Fewremarks regarding the morphologicalterminology used in this papermay be appropriate.The leavesof Leucobryaceaeconsist to the greaterpart of a multistratoseportion, containing inter alia the chlo- rocysts,which has traditionallybeen referredto as costa.Thus, the unistratosepart which is mostlycon- fined to the lowerparts of the leavesmay be termedlamina, or, as I havechosen, hyaline lamina, since it lackschlorocysts. As evidentfrom the discussionabove, the "familial"name Leucobyaceaeis hereapplied for the sake of convenienceonly, since this namecan be understoodto includeall mosseswith a "leucobryoidThabit. It is easier,then, for a personwho is not veryfamiliar with these mosses,to find the rightgenus and epi- thet for anygiven specimenbelonging to this group- whichin a phylogeneticalsense naturally is a non- group. I have also studiedthe Leucobgyaceaeof Western Melanesia, which is understoodto includeWest Irian,Papua New Guinea and the SolomonIslands; a forthcomingpaper (Enroth 1990, in prep.)will con- tain line drawingsof severalspecies of Leucobgyaceaealso known from Borneo. In Borneoseven generaand 23 speciesof Leucobgaceaeoccur. 16 namesare reduced to synonymy, i.e. Leucobiyumarfakianum C. Muell.ex Geh.,L. brassiiBartr., L. byssaceumC. Muell.exJohnson, L. cyathi- folium Dix., L. papuensePar., L. pentastichumDozy & Molk.,L. scalareC. Muell. ex Fleisch.,L. sericeum Broth.ex Geh.,L. subsanctumBroth., L. subscalareBroth. ex P.Vard., L. uncinatumFleisch., longifoliumLindb., 0. smaragdinumMitt., Schistomitriumlowii Mitt., S. nieuwenhuisiiFleisch. and S. subro- bustumBroth. Manyspecies have a wide geographicalrange and exhibitgreat morphological variability. However, I am convincedthat, in most cases,practically nothing is gainedby separatingsubspecific taxa, if the geo- graphicaland/or morphologicallimits between them cannot be clearlyindicated. Essentialbackground information, including the explanationof the Romannumerals refering to the collectingsites, is given in the first part of this series(Menzel 1988).

Key to the Bornean genera of Leucobtyaceae

1. Leaveswith a centralstereid band ...... Leucophanes - Leaveswithout a centralstereid band ...... 2 2. Chlorocystsin three layersin the upperparts of leaves ...... 3 - Chlorocystsin one layerin the upperparts of leaves ...... 4 3. Superficialchlorocyst layers exposed; leaf subula distinctlyscabrous ...... Exostratum - Superficialchlorocyst layers embedded under a hyalocystlayer; leaf subulasmooth ...... Arthrocormus 4. Chlorocystsmostly triangularin cross section in the upperparts of leaves ...... Octoblepharum - Chlorocystsmostly quadrangularin cross section throughoutleaves ...... 5 5. Adaxialhyalocysts near leaf apex multistratose;calyptra fimbriate at base ...... Schistomitrium - Adaxialhyalocysts near leaf apex unistratose;calyptra entire or lobed at base ...... 6 6. Chlorocystshypocentric throughout leaves; hyaline laminaonly up to 3(-4) cells wide; leaf apex smooth abaxially,usually piliferous or truncate,rarely apiculate; capsule erect, symmetrically cylin- drical,smooth ...... Cladopodanthus - Chlorocystscentric to hypercentricabove; hyaline laminafrom 3 to over 10 cells wide; leaf apex smooth or scabrousabaxially, mostly acute, apiculate, never piliferous or truncate;capsule inclined to cernuous,asymmetrical, sulcate ...... Leucobiyum Willdenowia18 - 1989 531

LeucobtyumHampe in Linnaea13: 42 (1839).

Leucobiyumis widelydistributed within the warmand temperateregions of the world,but the highest species diversityis encounteredin the tropics.In Borneo 8 species occur.

Key to the Borneanspecies of Leucobiyum

1. Matureleaves with distinct hyaline auricles...... L. sanctum - All leaveswithout auricles...... 2 2. Hyaline lamina in the lowerparts of leaveswide, consistingof 7-12 longitudinalcell rows . . 3 - Hyaline lamina in the lowerparts of leaves narrow,consisting of 2-5(-6) longitudinalcell rows ...... 3. Leavesup to 6 mm long; costalhyalocysts near leaf base mostly in 3 layersabaxially and in 2(-3) lay- ers adaxially...... L. neilgherrense - Leaveslonger (up to 15 or 16 mm);costal hyalocysts near leaf base mostly in 2 layersabaxially and in 1-2 layersadaxially ...... 4 4. Leavesup to about 10 mm long; adaxialcostal hyalocystsnear leaf base mostly in 1 layer ...... L. bowringii - Leavesup to 15(-16) mm long; adaxialcostal hyalocystsnear leaf base mostly in 2 layers ...... L. pulchrum 5. Plantsless than 1 cm high; leaves narrowlanceolate with abaxiallysmooth tips; hyaline lamina vanishingbelow leaf apex ...... L. chlorophyllosum - Plantsfrom 1 to well over 10 cm high (orlong); leaves lanceolate with abaxially more orless scabrous tips...... 6 6. 1-1.5 cm high; costal hyalocystsnear leaf base in (1-)2 layersabaxially and in 1(-2) layers adaxially...... L. stenophyllum - Plantsusually larger; costal hyalocysts near leaf basein 2-3(-4) layersabaxially and in 1-2(-3) layers adaxially...... 7 7. Plantsup to about2.5 cm high; leavesup to 3 mm long; superficialabaxial costal hyalocysts nearleaf base short, mostly quadrateto short rectangular,or irregularbut about as long as wide ...... L. aduncum - Plantsfrom about 2 towell over10 cm high (orlong);leaves 3-15 mm long; superficialabaxial costal hyalocystsnear leaf base mostlyrectangular, or irregularbut longerthan wide, few about as long as wide...... L.javense

Leucobiyumaduncum Dozy & Molk. in Miq., Pl.Jungh. 3: 319 (1854). Typus:Java,Junghuhn s.n. (Iso- H-BR!). = Leucobryumscalare C. Muell.ex Fleisch.,Musci Buitenzorg 1: 143 (1904).-Typus:Indonesia,Java,"Tji- pannas,Sindanglaya! an Felsen,100 m, immersteril in 9 Exemplaren",Fleischers.n. (not seen);"am Pa- pandajanbei Garoet!1300 m",Fleischers.n. (not seen);Singapore,"am Bukit-Timah! 100 m",Flrischers.n. (not seen);Philippines, Luzon, Micholitz 173 (Isosyn-H-BR!); "Sumbawa:Sambori", Warburg s.n. (not seen). = Leucobryumsubscalare Broth. ex P.Vard.in Rev.Gen. Bot. 29:292 (1917).-Typus: Indochina,"Plaine de Chu-Maia Thua-Liu,pied des arbrisseaux",Eberhardt 416-417 (not seen).- Note: In H-BR two speci- mens from Burma("Southern Shan States")are conserved under the name L. subscalare.They were col- lectedin 1899byW. Micholitz, and presumably described by Brotherusin a letterto P.de laVarde. Hence, the latter'sidea of L. subscalarewar obviously based on these specimens,although he did note cite them 532 Enroth:The bryophytesof Sabah(North Borneo)IV.

in the protologue;therefore I feel justifiedto synonymizeL. subscalarewith L. aduncumwithout having seen the types of the former. = Leucobryummicropentastichum C. Muell., Genera Musc. Frond. 81 (1900),nom. nud. - Originalcollec- tion: India,"Moulmain Hills", Stoliczka 4443 (H-BR!).

Ic.: Dozy&Molkenboer1854-1870: pl. 11;Cardot 1900: pl. 4, figs. 14a,b,fig. 15 (asL. scalare);Fleischer 1904: 142,fig. 18; Bartram1939: pl. 5, fig. 77 (asL. scalare);Johnson1964: 353, fig. 9k (asL. scalare),355, fig. 11; Gangulee1971: 429, fig. 201,430, fig. 202 (as L. scalare);Yamaguchi &Iwatsuki 1987: 483, pl. 6, figs. 1-22 (asL. scalare),485, pl. 7,figs. 1-20 (asL. scalare),487, pl. 8, figs.6,7 (asL. scalare),489, pl. 10, figs. 1-6 (as L. scalare).

Plantsup to about2.5 cm high,forming rather compact tufts. Leaves up to 3 mm long,nearly straight to falcate,from ovate baserather gradually narrowing into a semitubuloseacumen, tips usuallystrongly scabrousabaxially. Hyaline lamina narrow, near leaf base consistingmostly of 4-5 rowsof cells.Costal hyalocystsnear leaf base in 2-3 layersabaxially and in 1-2 layersadaxially, above in a singlelayer on both sides.Superficial abaxial hyalocysts near leaf base quadrate or irregularin shapebut mostlyabout as long as wide. Chlorocystshypercentric. Dioicous (pseudautoicous?).Female gametoecia lateral and pseudolateral.Seta mostly 1.5-1.6 mm, smooth.Capsule 1.2-1.3 mm long, mostlyhorizontal, strumose, sulcate when dry.Peristome about 0.35 mm high, otherwiseas in L. sanctum.Male plants not observed. L. aduncumlooks verymuch like a miniatureL.javense, but seems to be consistentlydistinguished from the latterby the quadrate,more or less not rectangular,superficial abaxial hyalocysts near the leaf base.Compared to L. stenophyllum,on the otherhand, L. aduncumhas wider and shorter,more crowded leavesand usuallythe costal hyalocystsnear the leaf base are in 2-3 layersabaxially and in 1-2 layers adaxially.In L. stenophyllum,these cells are in (1-)2 layersabaxially and mostly in 1 layeradaxially. Yamaguchi& Iwatsuki (1987) distinguished L. aduncumfrom L. scalareby the lengthof the perichae- tial leavesaround mature sporophytes relative to the length of the ordinaryleaves, but as faras I under- stand,this charactershows no consistencywhatsoever and has no taxonomicvalue. L. aduncumgrows most often as an epiphyteand on decayingwood, but is ratherfrequently encoun- teredon soil or humusas well.Ecological data from both Borneanand NewGuinean specimens suggest that it is restrictedto lowermontane forests,rarely occurring above 1500 m.

Geographicalrange: China; widely distributed from Nepal,India, and Sri Lankato Thailandand Viet- nam, Malaysia,Indonesia (Borneo: SAB, SAR, KTI, KSE), Philippines, New Guinea,Solomon Islands, New Caledonia,Vanuatu, Fiji, Society Islands,Australia (Queensland). Altitudinalrange (Borneo): 1050-1500 m. Specimensexamined: BRYOTROP:XI: 4331, 4334. -XIII: 3006. -XV: 3495. Additionalcollections: KSE: Bandjarmassin,Tamnalagang, com. Grebe(H-BR). -Without locality,Kor- thalss.n. (H-BR).

Leucobtyumbouringii Mitt. inJ. Proc.Linn. Soc. Bot. Suppl. 1: 26 (1859). Typus:India, "In mont. Khasia,reg. temp.",J. D. Hooker&T Thwaites1272 (Iso-H-BR!, H-SOL!), 1275; Sri Lanka,Gardner 1279; Hong Kong, Bowrings.n. - Leucobryumsericeum Broth. ex Geh.in Biblioth.Bot. 44: 26 (1898). Leucobryumbowringiivar. sericeum (Broth.ex Geh.)Dix. in Ann. Bryol.5: 24 (1932).-Typus: Indonesia, Borneo, Mt Paojem,1876, Teysmann 11140 (Holo- H-BR!). Willdenowia18 - 1989 533

Ic.: Geheeb 1898:pl. 4 (as L. sericeum);Cardot 1900: pl. 1, figs. la, b (as L. sericeum),pl. 4, fig. 16, pl. 6, figs.26a-c (asL. angustifolium);Cardot 1905: 97, fig. 4 (asL. confine);Bartram 1939: pl.5, fig.74;Johnson 1964:353, figs.9f, g, 354, fig. 10b; Gangulee1971: 419, fig. 195;Iwatsuki 1977: 361, figs.3o-s; Noguchi 1987: 239, fig. 96c.

Plantsmostly 1-2 cm high. Leavesup to 1 cm long, irregularlycurved to stronglyfalcate, obovate- lanceolate,upper parts flexuous, semitubular, tips smooth abaxially,apices acute, sometimes weakly den- tate;apical rhizoids common on leaves.Hyaline lamina wide below, consistingof several(up to 12)rows of linearto narrowrectangular cells with porosewalls. Costal hyalocysts at leaf basein 2 layersabaxially and in 1 layeradaxially, above in 1 layeron eitherside. Chlorocystshypercentric. Dioicous.Female gametoecia lateral and pseudolateral.Male plants and sporophytesnot observed. L. bowringiiis a wide-rangingand ratherhighly variable species, but easilydistinguished from most other membersof the genus by its broadhyaline laminanear the leaf base,which consistsof narrow, linearto long rectangularcells. This characteris also found in L. pulchrum;for the differencebetween these two species,see discussionunder the latter. L. sericeumis a form with stronglyfalcate leaves, giving the cushionsa nice "combed"appearance. However,in the copious materialI have studiedfrom Indonesia and adjacentregions, there are so many intermediatesbetween the "sericeum-type"and the "normaltype", that I haveconcluded not to recognize L. sericeumat any taxonomiclevel. TypicallyL. bowringiigrows on groundor humusin shadedhabitats, but it hasalso been collectedon treebases. Ecological data for the Borneanspecimens studied are practically lacking, but in New Guinea L. bowringiioccurs in lowermontane forestsbetween approximately1200-2000 meters.

Geographicalrange: Japan, China, the Himalayanregion, India, Sri Lanka; widely distributed in Indo- china,Malaysia, and Indonesia(Borneo: SAR, KBA), New Guinea,Philippines, Australia (Queensland), Vanuatu. Specimensexamined: BRYOTROP:none. Additionalcollections: KBA: see type cited above.- SAR:Mt Poe, Everett46 (H-BR).

Leucobryumchlorophyllosum C. Muell., Syn. Musc. Frond.2: 535 (1851). Typus:Indonesia, "Insula Sumbawa Archipelagi Indici", Zollinger 3370 (not seen).

Ic.: Dozy & Molkenboer1854-1870: pl. 10.

Plantsless than 1 cm high.Leaves oblanceolate-lanceolate, mostly 2.5-3 mm long, nearlystraight to slightlycurved, tips smooth abaxially,apices acute-acuminate; apical rhizoids quite common on leaves. Hyalinelamina narrow, vanishing below leaf apex.Costal hyalocysts at leafbase in 1(-2) layer(s)abaxial- ly and in 1 layeradaxially, above in 1 layeron eitherside. Chlorocysts hypercentric below, mostly centric above. Dioicous?Gametoecia and sporophytesnot observed(description based on NewGuineanandJava- nese specimensin H and H-BR). L. chlorophyllosumhas been recordedfrom Borneo only once, as"fo. minorFleisch." (Dixon 1916).It is a smallplant, confusable with L. stenophyllum;for the differencesbetween these two, see the discussion underthe latter.Because the specieswas not collectedduring the BRYOTROPexcursion, the description given above is based on specimensfrom Java and New Guinea. Accordingto Fleischer(1904), L. chlorophyllosumis autoicous. I have not seen sexualorgans in this 534 Enroth:The bryophytesof Sabah(North Borneo) IV.

species,but I stronglydoubt thatFleischer is correct.To the bestof my knowledge,all otherspecies of the genus aredioicous, and it is likelythat Fleischer confused dwarf males with "ordinary"male gametoecia. The ecologyofL. chlorophyllosumis not well known,but at leastin NewGuinea it seemsto be restrict- ed to lowlandrain forests, viz. belowapproximately 1000 m. Obviouslythis speciesmost often growsepi- phyticallybut at least occasionallyalso on decayingwood.

Geographicalrange: Sri Lanka,Malaysia, Indonesia (Borneo: SAB), Philippines, New Guinea. Altitudinalrange: not known for Borneo.In New Guinea 400-760 m. Specimensexamined: BRYOTROP:none.

Leucobryumjavense(Brid.) Mitt. in J. Proc.Linn. Soc., Bot. Suppl. 1: 25 (1859). -Sphagnumjavense Brid.,Muscol. Recent. 2(1): 27 (1798)(nom. inval.)et Bot. Zeitung(Regensburg) 1: 200 (1802).- Typus:Indonesia,Java, Commerson s.n. (not seen). ? = Dicranumcandidum Brid. ex P. Beauv.,Prodr. Aetheogam. 53 (1805) Leucobryumcandidum (P. Beauv.)Wils. in Hook. fil., Fl. Nov.-Zel.2: 64 (1854).- Typus:Australia (not seen). - Leucobgyumpentastichum Dozy & Molk.in Miq., Pl.Jungh.3: 319 (1854) Leucobtyumcandidum var. pentastichum(Dozy &Molk.) Dix. in New ZealandInst. Bull. 3: 97 (1923).- Typus:Indonesia,Java,Jung- huhns.n. (Isosyn-H-BR!). ?= Leucobiyumteysmannianum Dozy&Molk.,Bryol.Jav. 1: 17(1855).-Typus:Indonesia,Java,com.Teys- mann (not seen). ?= Leucobryumcandidum var. majusjaeg. in Ber.Thatigk. St. GallischenNaturwiss. Ges. 1871-72: 311 (1873).- Typus:Australia (not seen). = Leucobgumuncinatum Fleisch., Musci Buitenzorg 1: 149 (1904).- Typus:Indonesia, Borneo, Nieuwen- huis s.n. (Holo- BO, iso- H-BR!). = Leucobryumgyathifolium Dix. inJ.Linn.Soc.Bot.45:499 (1922).-Typus:Papua NewGuinea,"PortMo- resbyDistrict, Mt Durigolo",1916, Clark9 (Iso- H-BR!). = Leucobiyumbrassii Bartr. in Brittonia9: 36 (1957).- Typus:Papua New Guinea,"Milne Bay District, N slopesofMtDayman,2230 m,onground and lowertreetrunks in mossyforest",23.5.1953,Brass22466 (Holo- FH, iso- H!). - Leucobryumtranslucens C. Muell. in Par., Index Bryol. Suppl. 230 (1900), nom. nud. - Original collec- tion: PapuaNew Guinea,"Nova Guinea austro-or.brit. in mont. Yule",1895, Kowald1300 (H-BR!). = Leucobiyumpseudocandidum Broth. in Broth.& Wattsin Proc.Linn. Soc. New South Wales40: 366 (1915),nom. nud. - Originalcollections: Lord Howe Island,Watts 120, 148, 173,521, s.n., Whitelegge14 (all H-BR!).

Ic.: Dozy & Molkenboer1854-1870: pl. 14 (as L.falcatum),pl. 15 (as L. pentastichumand L. teysmannia- num);Cardot 1900: pl. 4, fig. 12,pl. 6, figs.23a, b (asL. conocladum),figs. 24a-d (asL. brachyphyllum);Th& riot 1921:222, fig. 1 (asL. candidum),223,fig. 2 (as,L.brachyphyllum); Bartram 1939: pl. 5, fig. 76 (asL. can- didum);Sainsbury 1955: 101, pl. 16, figs. 1-4 (as L. candidum);Johnson 1964: 353, fig. 9e (as L. cyathi- folium),9n, 354, fig. 10d (as L. candidum),10e (as L. cyathifolium);Gangulee 1971: 427, fig. 200; Scott, Stone &Rosser 1976: 161, pl. 26; Noguchi 1987:242, fig. 98;Yamaguchi & Iwatsuki 1987: 484, pl. 7 (asL. candidum),487, pl. 8, figs. 8, 9 (as L. candidum),489, pl. 10, figs. 7-9 (as L. candidum).

Plantshighly variable in size and habit, epiphytic,on decayingwood or on soil. Leaves3-15 mm long, sometimesdistinctly 5-seriate, lanceolate, tips scabrousabaxially. Hyaline lamina rathernarrow, up to 5(-6) cell rowswide nearleafbase. Costal hyalocysts at leafbase in 1-2(-3) layersadaxiallyand 2-3(-4) Willdenowia18 - 1989 535

layersabaxially, above in one layeron eitherside. Superficial abaxial hyalocysts nearleaf base mostlyrec- tangular,few quadrateor about as long as wide. Dioicous (pseudautoicous).Female gametoecia lateral. Seta 2.5-3.5 cm long, mostlyerect, smooth. Capsuleinclined to cernuous,1.5-1.8 mm long, strumose,sulcate. Peristome as in L. sanctum.Male plants about 1 mm high, anchored on leaf axils or elsewhereon the vegetativeplants. I am personallyconvinced that L. candidumand L. teysmannianumare conspecific with L.Javense, but since I have not seen the types of the two firstmentioned taxa, the questionmarks seem to be appropri- ate. Traditionally,the nameLeucobgyumjavense hasbeen usedonly forthe largerformsof this species,and L. candidumfor the smallerones. However, in this casea highly,or, should I say,almost unacceptably,var- iablespecies should obviously be recognized,but the variabilityconcerns superficial characters only. The cellularstructure of the leavesis essentiallythe samein the largestas well as the smallestforms.The large formswith leavesup to about 15 mm long seem to be geographicallymore restricted,occurring in the tropicalparts of Indonesiaand Australasia, and the Ryukyus.Still, there are no morphologicalgaps to be found what so ever,and subspecificdivision would not be appropriate. The smallforms of L.Javensemay be confusedwith the closelyrelated L. aduncum.However, as stated byYamaguchi& Iwatsuki (1987) - althoughthey speak of L. scalareand L. candidum,whichI considercon- specificwith L. aduncumand L.javense,respectively - the apparentlyconsistent difference is to be found in the superficialabaxial hyalocysts near the leaf base.In L. aduncumthey aremostly quadrate or at least aboutas long aswide, whereas in L.Iavensethey are mostly rectangular or at leastclearly longer than wide. L.javensehas a widealtitudinal range (see below), but NewGuinean and Bornean collections have gi- ven me the impressionthat it is most common in uppermontane forests, between approximately 1800- 2500 m.This speciesgrows on soil and asan epiphyte;ofthe 21 BRYOTROPspecimens14 were epigaeic and 9 epiphytic.

Geographicalrange: Widely distributed from India and SriLanka through SE-Asia and Australasia; also in the RyukyuArchipelago, New Caledonia,Tasmania and NewZealand. Borneo: SAB, SAR, KBA, KSE. Altitudinalrange (Borneo): 900-3400 m. Specimensexamined: BRYOTROP:IX: 4416. -XI: 4280,4310,4335,4336. -XII: 3005,3038. -XV: 3507 -XVII:3434,3464. - XIX: 3094, 3182, 3213. -XXI: 3237, 3244, 3319, 4861. - XXIII:3343, 3362. -XXXIV: 3839, 3841. Additionalcollections: SAB: Mount KinabaluTrip, Mount Bungal,on ground,9.- 10.12.1915, Mary StrongClemens 11226 (H-BR). - SAR:Matang Road, 8. 1911,Native collector 666 (H-BR).- KBA:Bukit Mulu, 1000 m, 2. 12. 1924, Winkler3074 (H-BR).-KSE: Bandjarmassin,Tamnalagang,com. C. Grebe s.n. (H-BR).

Leucobryumneilgherrense C. Muell. in Bot. Zeitung(Berlin) 12: 556 (1854). Typus:India, Neilgher, Perrottet s.n. (not seen).

Ic.: Dozy &Molkenboer 1854-1870: pl. 13 (asL. hollianum("Holleanum"); Cardot 1900: pl. 3, fig. 8 (asL. retractum),pl. 6, fig.28 (asL.japonicum); Warnstorf 1916: 81, fig. 17(asL. rhizophyllum);Bartram 1939: pl. 5, fig. 75;Johnson1964: 353, fig. 9a; Gangulee1971: 423, fig. 198 ("nilghiriense");Iwatsuki 1977: 360, fig. 2; Noguchi 1987: 237, fig. 95.

Plantsusually less than2.5 cm, but sometimesup to 4-5 cm high,growing in loose to compacttufts or cushions.Leaves up to 6.0 mm long, erectto spreading,ovate-lanceolate, upper parts (semi)tubulose, 536 Enroth:The bryophytesof Sabah(North Borneo) IV.

apicesabaxially smooth, acute-acuminate,often apiculate.Costal hyalocysts near leaf base mostlyin 3 layersabaxially and in 2(-3) layersadaxially, above in 1 layeron eitherside. Hyaline lamina nearleaf base wide,composed of 1-3(-4) rowsof linearcells followedby (4-)5-6(-8) rowsof shorter,mostly rectangu- lar or quadratecells. Dioicous?Gametoecia and sporophytesnot observed(description based on New Guineanmaterial in H). Accordingto Touw(1978), L. neilgherrensehas been recordedfrom Borneo only byHampe(1872). The speciesis somewhatless variable than many otherLeucobryum species and readilyrecognized by the wide hyalinelamina which consistsof distinctlywider cells than in L. bowringiiand L.pulchrum. In addition, near the leaf base the costal hyalocystsof L. neilgherrenseare in 3 layersabaxially and in 2(-3) layers adaxially,which servesto distinguishthe species from all other Borneanmembers of the genus. Naturally,the ecologyof L. neilgherrensein Borneo is not known.In New Guinea,however, the spe- cies has been collectedbetween 860-1860 m (Enroth1990, in prep.),whereit seemsto be most often en- counteredin lowermontane forests.The substratesinclude soil, rocks,tree roots,and decayingwood.

Geographicalrange: From India and SriLanka to China,Korea,Japan, and the Bonins;Indochina, Ma- laysia,Indonesia (Borneo: SAR), Philippines, New Guinea. Specimensexamined: BRYOTROP:none.

Leucob?yumpukhrum Broth. in Mitt. Inst.Allg. Bot. Hamburg7: 118 (1928) - Fig. 1. Typus:Indonesia. Borneo,"West-Borneo, aufdem BukitRaja, c. 1400m",21. 12.1924, Winkler3196(Ho- lo- H-BR!).

Plantsvery robust, several cm long, glossy.Leaves up to 15(-16)mm long,patent to spreading,some- times ratherweakly falcate, from ovate base narrowedinto a long, flexuous,semitubulose acumen, tips smooth abaxially,apices acute, sometimes apiculate, often weaklydentate. Hyaline lamina wide at leaf base,where mostly consisting of 10-12 rowsof linearto long rectangularcells with porosewalls. Costal hyalocystsnear leaf basein (1-)2 layersboth adaxiallyand abaxially, above in 1 layeron eitherside. Chlo- rocystshypocentric to centric below, hypercentricabove. Dioicous. Femalegametoecia lateral. Perichaetial leaves up to about 3.5 mm long, from broadly ovate base suddenlynarrowed into a lanceolateacumen. Sporophytes and male plants not observed. In termsof the leaf structure(Fig. 1),L. pulchrumseems to be ratherclosely related to L. bowringii. However,the formeris distinctlymore robust,has muchlonger and at basebroader leaves than anyspe- cimens of the latterspecies I have seen. The largestforms of L.javensemay also be confusedwith L. pulchrum,but the formeris neverquite as glossy,its hyalinelamina is clearlynarrower (only up to 6 cells wide),it hasthe costalhyalocysts near the leafbase partly in 3 layers,and the leaf tips areusually strongly scabrous. The total numberof collectionsof L. pulchrumis low, and hence its ecologyremains insufficiently known.Only two specimenswere collected during the BRYOTROPexpedition. Theywere encountered in lowermontane forests,one was growingepigaeically, the other epiphytically.

Geographicalrange: Malaysia (Pahang), Borneo (SAB,KBA), Sumatra. Altitudinalrange (Borneo): 1850-1950 m. Specimensexamined: BRYOTROP:XIXb: 4718, 4744. Additionalcollections: see type cited above. Willdenowia18 - 1989 537

a

2 mm 100Pm

K,~~~~~~~~~~~~~j

100Pm C i

~~ctct~~~~~

Fig.1. Leucobryum pulchrum Broth. (from the holotype).- a: Leaves;b: Leafmargin and hyaline lamina at 1/4 leaf lengthfrom base; c: Crosssection from leaf margin to medianline nearleaf base. 538 Enroth:The bryophytesof Sabah(North Borneo) IV.

Leucobryumsanctum (Brid.) Hampe in Linnaea13: 42 (1839). - Dicranumglaucum Hedw. var. sanctum Brid., Bryol. Univ. 1: 811 (1827). Typus:Indonesia, Java, "in tumulis sanctisBaduorum in insulaJava", Blume s.n. (not seen). = Leucobryumpapuense Par., Index Bryol. 752 (1897).- Lectotypus(designated here): New Guinea,"Neu Guinea",23. 6. 1875,Naumann s.n. (H-BR!). = Leucobiyumarfakianum C. Muell.ex Geh.in Biblioth.Bot. 44: 5 (1898).-Lectotypus(designated here): New Guinea,"Nova Guinea,Mt Arfakad Hatam,5-7000"', 7. 1875,Beccari 163 (H-BR!). = Leucobiyumsubsanctum Broth. in Philipp.J.Sci. 2: 339 (1907).-Typus: Philippines, Mindoro,"Mt Hal- con, on cliffsat 1970m. alt.",1906, ElmerD. Merrill 6161 (Lecto-H-BR!);Philippines,"Luzon of Bataan, summitof Mt Mariveles",1903, ElmerD. Merrill3540, 3549; Mindoro,"Mt Halcon, on prostratelogs at 1350 m. alt.",1906, ElmerD. Merrill6208 (all syn- H-BR!). = Leucobyumbrevisetum C. Muell.in Par.,Index Bryol. Suppl. 228 (1900),nom. nud. - Originalcollec- tion: New Guinea,"Nova Guinea austro-orient. in montosis Moroka,1300 m, distr.Moresby", 7. 1893, Loria 743(H-BR!).

Ic.:Dozy&Molkenboer 1854-1870: pl. 12;Geheeb 1898:p1. 3 (asL. arfakianum);Cardot 1900: pl. 1,figs. 2a-d, figs.3a-d (asL. vesiculosum);Brotherus 1901-05: 346, figs.205; Brotherus1924: 223, fig. 178;Bar- tram 1939:pl. 5, fig. 72, fig. 73 (as L. subsanctum);Johnson1964: 353, fig. 9c (asL. subsanctum),fig. 9d; Gangulee1971: 417, fig. 194.

Plantslax, severalcentimeters long, growingin ratherloose mats or tufts mostlyon tree basesand roots.Leaves erect to spreading,ovate-lanceolate, up to 6(-7) mm long, basesof matureleaves with dis- tinct hyalineauricles, tip more or less scabrousdorsally, apex obtuse to acute,mostly apiculate. Hyaline laminanear leaf base3-4 cellswide. Costal hyalocysts in one layeron both sides of the chlorocystlayer, at base occasionallyabaxially 2(-3) layered. Dioicous (pseudautoicous).Female gametoecia lateral, bud-shaped. Seta about 2 cm, smooth, slen- der.Capsule about 1.5 mm long, inclinedto cernuous,asymmetric and curved,strumose, sulcate when dry.Peristome teeth 16, about 0.6 mm high, articulateand weaklylongitudinally striolate, papillose throughout,above divided into two lanceolate-subulate,knotted segments.Male plants dwarf-like, about 1 mm high, mostly attachedto the leaves of the gametophytes. Althoughsome gametophyticcharacters of L. sanctumshow ratherstrong variability, this speciesis alwayseasily recognized by the hyalineauricles at the basesof matureleaves. In young leavesnear the tips of the stems and branchesthe auriclesmay be very small or even absent. L. sanctumis primarilyan epiphyte,favoring tree roots,bases and lowertrunks as substrates,but it also growson decayingwood and soil.The BRYOTROP-specimens were collected in lowlandand lower montane rain forests,but in New Guinea the altitudinalrange extends up to about 2500 m.

Geographicalrange: NE-India, Indochina,Malaysia, widely distributedin Indonesia(Borneo: SAB, SAR,KBA, KSE, KTI), Philippines, New Guinea,Carolines, Fiji, Samoa, Australia (Queensland). Altitudinalrange (Borneo): 20-1700 m. Specimensexamined: BRYOTROP:Ia: 4549, 4553, 4570, 4571. - IX: 4406. - XVIIa:4628, 4774. Additionalcollections: SAB: Sandakanand vicinity, 1920,Ramos 2158, 2162 (H, H-BR). - SAR:Mt Poeh,8. 1892,Everett39 (H-BR). -KBA: Pontianak,Lohaban,7. 1894,Ledru2372 (H-BR).-KSE:"Prope Martapoera",Korthals s.n. (H-BR);Boekit Milik, 23. 9. 1898,Amd]ahs.n. (Nieuwenhuis-Expedition)(B).

Leucobiyumstenophyllum Besch. in Ann. Sci. Nat. Bot. ser.5, 18: 204 (1873). Typus:New Caledonia,Balansa 2531 (Holo- BM, iso- H-BR!). Willdenowia18 - 1989 539

= Leucobiyumbyssaceum C. Muell.exJohnson in Gard.Bull. Straits Settlem. 20: 334 (1964).-Typus: Pa- pua New Guinea,"Nova Guinea austro-orient. secus flumenS.Josephi, distr. Moresby", 11. 1892,Loria determinavitC. Muellersub) 703 (Lecto-BM, isolecto-H-BR!).

Ic.:Cardot 1900: pl. 5,figs.22a-d;Johnson1964: 353,fig. 8 (asL.bssaceum),354,fig.9h (as L. byssaceum), 9j,355, fig. 10c.-Yamaguchi &Iwatsuki 1987: 480, pl. 4,481,pl. 5,487,pl.8, figs.4,5, 488,pl. 10,figs. 6-9.

Plants1-1.5 cm high. Leavesup to about4 mm long, falcate,narrow obovate-lanceolate, tips more or less stronglyscabrous abaxially, apices mostly apiculate. Hyaline lamina narrow, at base4-5 cells,at tip only one cell wide.Costal hyalocysts near leaf base in (1-)2 layersabaxially, mostly in one layeradaxially, above in one layeron eitherside. Chlorocystshypercentric. Dioicous (pseudautoicous?).Female gametoecia pseudolateral. Seta 2-2.5 cm long, smooth. Cap- sule 0.8-0.9 mm long, horizontalto cernuous,asymmetric, strumose, sulcate. Peristome about 0.35 mm high, otherwiseas in L. sanctum. L. stenophyllumis newly recorded for Borneo.As a small species,it may be, and probablyhas been, confusedwith L. chlorophyllosum,but the formerhas the leaftips moreor less scabrousabaxially, whereas in the latterthey arealways smooth. In addition,the leavesof L. chlorophyllosumare usuallynot falcate and morenarrow than those of L. stenophyllum.L. aduncum, on the otherhand, is muchmore robust with morecrowded and widerleaves, and the abaxialcostal hyalocysts near the leafbase are partly in 3 layers. In New GuineaL. stenophyllumappears to be a strictepiphyte; the two BorneanBRYOTROP spe- cimens were also growingepiphytically in lower montane forests.

Geographicalrange: Borneo (SAB),New Guinea,Solomon Islands,New Caledonia,New Hebrides. Altitudinalrange (Borneo): 1050-1250 m. Specimensexamined: BRYOTROP:XI: 4346. - XII: 4672.

SchistomitriumDozy & Molk., Musci Frond.Ined. Archip. Ind. 3: 67 (1846).

Schistomitriumis closely relatedto Cladopodanthus,but distinctin the multistratoseadaxial hyalo- cystsnear the leaf apicesand the fimbriatebase of the calyptra.These two generaalso sharea rathernar- rowgeographical distribution, both being confinedto the Malaysian- Indonesian- Australasianregion. The gametophyticcharacters of Schistomitriumand Cladopodanthusrather closely resemble those of Leucobryum,but the sporophytesare completelydifferent. The capsulesof the two formergenera are erect,symmetrically cylindrical, and smooth on the surface,and the peristometeeth, 16 in number,are not dividedabove into two segments.Hence, I thinkthat the phylogeneticas well asgeographical origins of Schistomitriumand Cladopodanthusare differentfrom those of the widely distributedLeucobryum. The total numberof collectionsof any speciesof Schistomitriumand Cladopodanthusis not too high. However,this probablydoes not mean that these species are rarethroughout their range, but rather reflectsthe fact thatthey often growhigh in treecrowns and aretherefore not easilycollected (D. Norris, pers.comm.). For example, many new Guineanspecimens were collected on fallentwigs or branchesof trees. Touw's(1978) list of Borneanmosses includes4 speciesand one nomen nudum of Schistomitrium, but I recognizeonly 3 species,and, moreover,one of them with strong reservationsonly.

Key to the Borneanspecies of Schistomitrium

1. Hyaline lamina below consistingof 10-12 cell rows...... S. robustum 540 Enroth:The bryophytesof Sabah(North Bomeo) IV.

- Hyaline lamina below consistingof 3-7 cell rows ...... 2 2. Plantsup to 3 cm high; leaves up to 4.0 x 1.0 mm, usuallyfalcate ...... S. apiculatum - Plantsup to 4-5 cm high; leaves up to 6.5 x 1.8 mm, straightor falcate...... S. mucronfoium

Schistomitniumapiculatum (Dozy & Molk.) Dozy & Molk., Musci Frond.Ined. Archip.Ind. 68 (1846). -Syrrhopodon apiculatusDozy & Molk. in Ann. Sci. Nat. Bot., ser.3, 2: 315 (1844). Typus:Indonesia, "Java, Sumatra" (not seen).

Ic.: Dozy & Molkenboer1845-1848: pl. 24, 25; Bartram1939: pl. 5, fig. 68;Johnson 1964:347, fig. 2h, 348, fig. 3.

Plantsup to about3 cm high, growingin compacttufts. Leaves up to 4.0 x 1.0 mm, nearlyoblong to lanceolate,apices acute to hood-shapedand rounded,apiculate. Hyaline laminabelow 3-6 cell rows wide. Leaf marginoften serrulatearound midleaf, entire elsewhere. Dioicous.Female gametoecia terminal (pseudolateral). Seta up to 8 mm long, smooth.Capsule 1.6- 1.8 mm long, erect, cylindrical.Calyptra 2.5-3 mm, fimbriateat base. Male plants not observed. S. apiculatumis, to saythe least,very closely related to the variableS. mucronifolium,but distinguished fromit by the usuallysomewhat smaller size of the leaves,as well asthe plantsthemselves. I havenot seen the specimenof S. apiculatumvar. copelandii (Broth.) Bartr. that Dixon (1935)reported from Borneo (as S. copelandiiBroth.), but since this varietyis understoodto possesslonger leaves than is usualin S. apicula- tum,it seemspossible that Dixon's specimen is bestplaced in S. mucronifolium.It should be stressedthat I recognizeS. apiculatumwith greatreservations; it is possiblyconspecific with S. mucronifolium. The ecologyof S. apiculatumis not well known,but it appearsto be a strictepiphyte, as arethe other species of the genus. It has been collected in lowland and lowermontane forests.

Geographicalrange: Malaysia, Borneo (SAB,SAR, KBA, KTI), Java, Sumatra, Philippines. Altitudinalrange (Borneo): 500-1900 m. Specimensexamined: BRYOTROP:XV: 3547. - XVII:3468. - XIX: 3099, 3215. Additionalcollections: SAR: Mt Poe, 4500-5000', 1924,Mjdberg s.n. (H-BR). - KBA:Bukit Mehipit, c. 500 m, 8. and 10. 12. 1924, Winklers.n. (H-BR).

Schistomitriummucronifolium (C. Muell.) Fleisch.,Musci Buitenzorg1: 161 (1904). -Leucobryum mucronifoliumC. Muell., Syn. Musc. Frond.2: 536 (1851). Typus:Indonesia,Java, Blume s.n. (Holo- FH!). = Schistomitriumlowii Mitt. inJ. Linn.Soc., Bot.22: 302 (1886).-Typus: Malaya, Borneo, Sarawak, Mt Ki- nabalu,Low s.n. (Holo- NY!). = SchistomitriumnieuwenhuisiiFleisch.,Musci Buitenzorg 1: 161(1904).-Typus: Indonesia,Bomeo,Kali- mantanTimur, "auf Baumzweigen Bockit Milie im InnerenBorneos", Nieuwenhuis s.n. (Holo- BO, iso- FH!). = Schistomitriumsubrobustum Broth. in Leafl.Philipp. Bot. 6: 1976(1913). - Typus:Philippines, Minda- nao, Cabadbaran(Mt Urdameta),Province of Agusan, 10. 1912, Elmer14111 (Holo- H-BR!). = Schistomitriumcornutum Fleisch. ex Broth.in Mitt.Inst. Allg. Bot. Hamburg 7: 119 (1928),nom. nud.- Originalcollections: Indonesia, Borneo,"Bukit Mulu, c. 700 m", 3. 12. 1924, Winkler3O79and "Bukit Ti- lung,c. 500 m", 8.2. 1925,Winkler3310(duplicates in H-BR!).-Note: Thename Schistomitrium cornutum Willdenowia18 - 1989 541

is not includedirnWijk, Margadant & Florschutz (1967), but to the best of my knowledgethe namewas first published by Brotherus(1928).

Ic.: Fleischer1904: 162,fig. 21; Fleischer1913: pl. 3 (asS. nieuwenhuisii);Bartram 1939: pl. 5, fig.67 (asS. subrobustum),fig. 69 (asS. nieuwenhuisii);Johnson1964: 346, figs. l e-g, 347,fig. 2a (asS. nieuwenhuisiz),fig. 2c (as S. subrobustum),figs. 2d (as S. lowif),fig. 2e, 349, fig. 4 (as S. nieuwenhuisii).

Plantsfrom 1 to 4 or 5 cm high.Leaves variable in length/widthratio but up to 6.5 x 1.8 mm, erect and straightto stronglyfalcate, nearly oblong to lanceolate,upper parts mostly (semi)tubulose,apices acuteto roundedand hood-shaped,apiculate. Hyaline lamina below up to 6(-7) cellswide. Leaf margin serrulatearound midleaf or entire throughout. Dioicous (pseudautoicous).Female gametoecia terminal (pseudolateral).Seta 6-8 mm long, smooth. Capsuleabout 1.8 mm long, erect,cylindrical. Calyptra 3 mm long, fimbriateat base.Male plants 1.5-2.0 mm high, anchoredby rhizoidson the leavesof femaleplants (not observedin Bornean specimens). Thisspecies is highlyvariablein size andhabit.The name S. nieuwenhuisiihasbeen usedforthe small forms,and the namesS. lowiiand S. subrobustumfor the largerones which often have erectand straight leaves.Two additional characters used in distinguishingthe "species"mentioned above are the widthof the hyalinelamina and the shapeof the leafapices. However, the variationof thesecharacters is alsocon- tinuousand inconsistent,and in my opinion the differentforms do not warranttaxonomic recognition. Forthe differencesbetween S. mucronifoliumand S. apiculatum,see the specifickey and the discus- sion under the latterspecies. S. mucronifoliumis an epiphyte,collected in Borneo in lowland and lowermontane forests.

Geographicalrange: Malaysia, Borneo (SAB, KBA, KTI),Java, Sumatra, Philippines, New Guinea,Solo- mon Islands. Altitudinalrange (Borneo): 500-1900 m. Specimensexamined: BRYOTROP:XVIIa: 4631. - XIX:3126, 3189. Additionalcollections: SAB: see type cited above;Mt Kinabalu,Marai Parai Spur, 1.-4.12. 1915,Magy StrongClemens s.n. (H-BR). - KBA: see materialcited above.- KTI:see materialcited above.

Scbistomitriumrobustum Dozy & Molk., Bryol.Jav. 1: 21 (1855). Typus:Indonesia,Java, com. Teysmann (not seen);"in m. Gegerbintangprope Toegoe altit. 4000'", Kurz s.n. (Syn-L!; this specimenis cited in the 'Addenda"section on p. 222 of Dozy & Molkenboer1854- 1870).

Ic.: Dozy & Molkenboer1854-1870: pl. 17;Johnson 1964: 346, figs. la-d, 347,fig. 2f.

Plantsrobust, up to about 5 cm high. Leavesup to 6.5(-7.0)mm long, oblong to ovate-lanceolate, apices mostly cucullate,hood-shaped, apiculate. Hyaline lamina below consistingof 10-12 cell rows. Leaf marginusually entire throughout. Dioicous (pseudautoicous?).Female gametoecia terminal (pseudolateral).Seta 6-9 mm long, smooth.Capsule about 2 mm long, erect,cylindrical. Calyptra 3 mm long,fimbriate at base.Male plants not observed(description based on Javanesematerial in H-BR and H-SOL). S. robustumwas recordedfrom Borneoby Dixon (1935).I have not seen that specimen,or any Bor- nean specimensof the species. 542 Enroth:The bryophytesof Sabah(North Borneo)IV.

Thewide (10-12 cells)hyaline lamina on the lowerparts of the leavesmakes S. robustuman easilyre- cognizedspecies, although the most robustforms of S. mucronifoliumapproach it in size and habit.The ecology of S. robustumis verybadly known, but Fleischer(1904) cites specimenswhich were growing on rottenwood at 1300 and 2500 m altitude.Thus, the species appearsto occur also in uppermontane forests,at least in Java.

Geographicalrange: Borneo (KBA),Java. Altitudinalrange: not known for Borneo; in Java 1300-2500 m. Specimensexamined: BRYOTROP:none.

CladopodanthusDozy & Molk., Musci Frond.Ined. Archip.Ind. 3: 79 (1846).

Cladopodanthusis distinguished from Schistomitriummainly by the entireor lobed (not fimbriate) base of the calyptraand the unistratoseadaxial hyalocysts near the leaf apices.Apart from the calyptra, the sporophytesof these generaare nearlyidentical. I havedecided to acceptfour species of CladopodanthusforBorneo, although two of them seemto be more or less dubious.

Key to the Borneanspecies of Cladopodanthus

1. Leaf apices truncate,usually apiculate, never piliferous ...... C muticus - Leaf apices apiculateor piliferous,not truncate...... 2 2. Leaf apices apiculate,never piliferous ...... C microcarpus - Leaf apices mostly piliferous ...... 3 3. Plantsjulaceous; leaf apices mostly rounded,sometimes obtuse, hair-pointsup to about 30pm long ...... C.speciosus - Plantsnot julaceous;leaf apicesmostly acute, sometimes obtuse, hair-points not rarelyover 1 mm long ...... C. pilifer

Cladopodanthusmicrocarpus Dix. in J. Linn. Soc. Bot. 50: 74 (1935). Typus:Borneo, Sarawak, Ulu Koyan,c. 800 m, on fallentree trunk,22. 9. 1932,Richards 2034bis (Holo- BM!).

Ic.: Dixon 1935: pl. 1, fig. 5;Johnson 1964: 351, figs. 7e, f.

Plantsup to about2 cm high, growingin dense tufts.Leaves up to 2.5 mm long, oblong-lanceolate, tips (semi)tubulose,apex mostly acute, apiculate;leaf marginsentire below, usuallyserrulate at tips. Hyaline lamina 2-3 cells wide. Dioicous. Femalegametoecia terminal. Seta 4-5 mm long, smooth. Capsuleabout 0.5 mm long, erect,cylindrical. Calyptra cucullate, entire or weaklylobed at base. Male plants not observed. Thisis a peculiarplant, in habitalmost exactly like Schistomitriumapiculatum, but the structureof the leaf apicesand the non-fimbriatecalyptra base are of the Cladopodanthus-type.C. microcarpus is the only Borneanspecies to possess mostly acute, stronglyapiculate leaf apices,similar to those of the (lower) leaves of the New Guinea endemic C. heterophyllus(Fleisch.) Bartr., which is, however,conspicuously Willdenowia18 - 1989 543

more robust.In fact, C. microcarpusappears to be justa "copy"of C.heterophyllus, just strongly reduced in size. In generalterms, size is not a veryreliable diagnostic character in Schistomitriumor Cladopodanthus, - orLeucobiyum, for that matter-, but in this particularcase I find it appropriateto recognizeC. microcarpus as an independenttaxon. C. microcarpusis only knownfrom the type specimen,which was collected at ca. 800 m altitudeon a fallen tree trunk.

Geographicalrange: Borneo (SAR). Altitudinalrange: "c. 800 m" (Dixon 1935). Specimensexamined: BRYOTROP:none. Additionalcollections: see type cited above.

Cladopodanthusmuticus Broth. in Philipp.J. Sci. 31: 279 (1926). Typus:Philippines, Panay, Capiz Province,Mt Salibongbong,on old tree, 6. 1919,Martelino & Edano (Bur.Sc.) 35787 (Holo- H-BR!, iso- BM!). = Cladopodanthustruncatus Dix. inJ. Linn.Soc. Bot. 50: 73 (1935).-Typus: Borneo, Sarawak, Marudi, un- der300 m, on rottentrunk and earthin 'heathforest', 1 1. 1932,Richards 2675 (Holo-BM!). - cf. Bartram 1939.

Ic.: Dixon 1935: pl. 1, fig. 6; Bartram1939: pl. 5, fig. 71;Johnson 1964: 351, fig. 7g.

Plantsup to about 4 cm high, julaceous,growing in dense tufts.Leaves very crowded, mostly 2.0- 2.5 mm long, oblong, apices truncate,often apiculate.Hyaline lamina 2-3 cells wide. Dioicous.Female gametoecia terminal. Seta 5-6 mm long, smooth.Capsule 1.1-1.4 mm long, erect, cylindrical.Calyptra cucullate, lobed at base. Male plants not observed. C. muticusis very easily recognized at first sight by the tightly julaceous habit and the truncate leaf apices.It is interesting,however, that Fleischer(1912) described and illustratedsimilar apices for the up- perleaves of C. heterophyllus;the lowerleaves of C.heterophyllus have mostly acute apices, similar to those of C. microcarpusdescribed and discussedabove. In all the New Guineanspecimens of C. heterophyllusI havestudied the heterophyllyisnotveryclear,and the leaveshave mostlyacute apices. Still,I feel thatthe relationshipsof C. muticusand C. heterophyllusrequire further critical study. C. muticusis obviouslyquite a rarespecies in Borneo;it wasnot collectedduring the BRYOTROPex- pedition.The type of C. truncatuswas collected below 300 m on a rotten tree trunkand soil.

Geographicalrange: Borneo (SAR),Philippines. Altitudinal range(Borneo): "below 300 m" (Dixon 1935). Specimensexamined: BRYOTROP:none. Additionalcollections: SAE: see type cited above. - Borneo,s. loc., nativecollector 1203 (H).

Cladopodanthuspilifer Dozy & Molk., Musci Frond.Ined. Archip.Ind. 3: 80 (1846). Typus:Indonesia, "Java, ad arborumcorticem in monte Gede",Zippelius s.n. (not seen).

Ic.: Dozy &Molkenboer 1845-1848: pl. 28; Cardot1900: pl. 6, figs.3 1a-d; Fleischer1904: 154,fig. 19; Brotherus1901-05: 345, fig.204; Brotherus1924: 222, fig. 177;Johnson1964: 351, figs.7a, b, 352,fig. 8. 544 Enroth:The bryophytesof Sabah(North Borneo)IV.

Plantsrather robust, up to about 10 cm long, often decumbent.Leaves 3-4(-5) mm long, narrowly oblong to lanceolate,semitubulose above, piliferous, apices mostly acute, sometimes obtuse, hair-points not rarelyover 1 mm long. Hyaline lamina2-3 cells wide. Dioicous. Femalegametoecia terminal (pseudolateral). Seta up to 8-9 mm long, smooth. Capsule 1.4-1.8 mm long, erect,cylindrical. Calyptra 2.3 mm long, cucullate,lobed at base.Male plants not ob- served(description based on Javaneseand New Guineanspecimens in H and H-BR). C. pilifercan be confusedwith C. speciosusonly; the differencesare discussedunder the latter. C. piliferwas reportedfrom Mt Kinabaluby Bartram(1936), but not collected there duringthe BRYOTROP-expedition.I have not seen any specimensof the speciesfrom Borneo, and the ecological datafor Borneanspecimens are meagerin the literature.Bartram (1936) reportsC. pilifer from approxi- mately1200-1350 m altitudeon treeand twigs.In New Guineathe specieshas been collectedbetween 1100-1700 m altitudeon bushes and trees (Enroth 1990, in prep.).

Geographicalrange: Malaysia, Borneo (KTI,SAB),Java, New Guinea. Altitudinalrange (Borneo): approximately 1200-1350 m (Bartram1936). Specimensexamined: BRYOTROP:none.

Cladopodanthusspeciosus (Dozy & Molk.) Fleisch.,Musci Buitenzorg1: 156 (1904). -Syrrhopodon speciosusDozy & Molk. in Ann. Sci. Nat. Bot., ser.3, 2: 315 (1844). Typus:Indonesia, "Java, in monte Gede ad arborumcorticem", Zippelius s.n. (not seen).

Ic.:Dozy&Molkenboer 1845-1848: pl.26 (asSpirula speciosa); Fleischer 1904: 157,fig.20; Bartram 1939: pl. 5, fig. 70; Johnson 1964: 351, figs. 7c, d, 352, fig. 8.

Plantsseveral cm high, julaceous,growing in quite compacttufts. Leaves very crowded, appressed, oblong,3(-4) mm long,piliferous, apices mostly rounded, sometimes obtuse, rarely acute, hair-points up to about 30 pm long. Hyaline lamina2-3 cells wide. Dioicous.Female gametoecia terminal (pseudolateral). Seta about 6 mm long, smooth.Capsule 1.6- 2.0 mm long, erect,cylindrical. Calyptra cucullate, lobed at base.Male plants not observed(description based on Javaneseand New Guinean specimensin H, H-BR, and H-SOL). In general,C. speciosusis distinguishedfrom C.pilifer by the julaceoushabit and roundedor obtuse leaf apiceswith shorter(only up to about 30 pm long) hair-point-s.However, there are specimens where some leaveshave acute tips and alsosomewhat longer hair-points, and it is possiblethat we arefaced here with a single specieswhich sometimesproduces shorter, sometimes longer leaves and hair-points. Bartram(1936) cited two specimensof C. speciosusfrom the Mt Kinabaluregion, collected at 1500 and 1800 m altituderespectively. One specimenwas growingon rocks,which is ratheran exceptional substratefor Cladopodanthus.

Geographicalrange: Borneo (SAB, SAR,KTI),Java, Sulawesi, Philippines; also New Guinea (Enroth 1990, in prep.). Altitudinalrange (Borneo): approximately 1500-1800 m (Bartram1936). In New Guineacollected at 500 m. Specimensexamined: BRYOTROP:none.

LeucophanesBrid., Bryol. Univ. 1: 763 (1826).

Leucophanesis unique within the "leucobryoid"group in havinga distinctband of centralstereids in Willdenowia18 - 1989 545

the leaves.One could be temptedto referto these stereidsas costa,but this would be inconsistentwith the traditionaluse of the word costa in this group. In addition,Leucophanes possesses marginally on the total lengthof the leaveslong, stereid-likecells, a characteralso found in the othergeneraof CalymperaceaesubfamilyLeucophanoideae,with the exception of Octoblepharum. Throughthe kindnessof Dr.D. H. VittI have in my possessiona copy of the asyet unpublishedmo- nographof Leucophanesby SalazarAllen (1986).She accepted12 speciesof Leucophanesfor the whole world,which means that a considerablenumber of nameshas been reducedto synonymy.Taking the high degreeof variabilityexhibited by manyspecies of the genus into consideration,this is not surpris- ing, and I mostlyfind SalazarAllen's results highly acceptable. All the acceptedspecies were illustrated and mapped,but since the draftof herpaper in my possessionis probablynot the finalversion and the page numberswill be changed,I prefernot to cite herexcellent drawings and SEM-photographs.How- ever,I thinkit is reasonableto includeSalazarAllen's (1986) new synonymshere, if they havebeen used for Borneancollections, even if theirtype specimensare not Bornean.Salazar Allen also lectotypified most species of Leucophanes,and the lectotypescited here were designatedby her. In Borneo four species of Leucophanesoccur.

Key to the Borneanspecies of Leucophanes

1. Abaxialcostal hyalocystsnear leaf base in 2-3 layers...... L. glaucum - Costal hyalocystsnear leaf base in 1 layeron eitherside of the chlorocystlayer ...... 2 2. Costal hyalocystsin upperparts of leaves partlyin 2 layerson eitherside of the chlorocystlayer ...... L. angustifolium - Costal hyalocystsin 1 layeron eitherside of the chlorocystlayer throughout the leaves..... 3 3. Leavesup to about 4 mm long, leaf base obovate,sheathing, upper parts usually squarrose, keeled ...... L. candidum - Leavesup to 6.5 mm long, leaf baseoblanceolate, not sheathing,upper parts not squarrose,plane or weaklykeeled ...... L. octoblepharioides

Leucophanesangustifolium Ren. & Card.in Rev.Bot. Bull. Mens. 9: 395 (1891). Typus:Reunion Island("Hab. Bourbon"), Rodriguez 74c (Lecto- PC, isolecto-PC, S).

Ic.: Cardot1897: pl. 1A;Cardot 1900: pl. 11,figs. 56a-e (asL.prasiophyllum), figs. 58a-c; Onraedt1976: 373, fig. 15; Whittier1976: 99, figs. 25J-L (as L. prasiophyllum),figs. 25M-O (as L. tahiticum).

Plantsup to about2 cm high,growing in ratherloose tufts.Leaves up to 7.5(-8.6)mm long, erectto patent,straight or weaklycurved, narrowly lanceolate to linear,keeled, apices mostly acute, dentate, api- cal gemmaecommon on leaves.Costal hyalocysts near leaf base in one layerbothadaxially and abaxially, above partlyin two layerson eitherside. Dioicous. Femalegametoecia terminal (pseudolateral). Sporophytes and male plantsnot observed. This speciesmay be confusedwith L. octoblepharioides.In general, L. angustifoliumhas longerand at leastrelatively narrower leaves, but for a positiveidentification cross sections through the upperparts of leavesare required.These will revealthe costalhyalocysts partly in two layerson eithersideof the chloro- cyst layer;in L. octoblepharioidesthe costalhyalocysts are always in justone layerboth adaxiallyand aba- xially. L. angustifoliumwas first recorded forBorneo by SalazarAllen(1 986). In hermapthere are two dotsin Borneo,and they appearto be placed in the provincesof KalimantanTimur and KalimantanSelatan. 546 Enroth:The bryophytesof Sabah(North Borneo)IV.

However,she does not cite the specimensstudied, and therefore I haveadded question marks in the para- graphon distribution. It should be mentioned here that the name L. angustifoliumhas previouslybeen used for African plants only, and the name underwhich this species has been known in the Indonesian- Australasian regionis L. massartiiRen. & Card., which was reducedto synonymyby SalazarAllen(1986). The BRYO- TROP materialincludes one specimen of the species. L. angustifoliumoccurs in the lowlandand lowermontane forests up to about 2000 m. It growson trees,decaying wood, and rocks.

Geographicalrange (from SalazarAllen 1986):Cameroon, Ghana, Tanzania, Comores (Mayotte), La Reunion,Seychelles (Mahe), India, Taiwan, Vietnam, Thailand, Malaysia, Sumatra,Java, Borneo (SAB, KBA,KTI?, KSE?), Halmahera, New Guinea,Australia (Queensland), Samoa, Society Islands. Altitudinalrange (Borneo): 1250 m. Specimensexamined: BRYOTROP:XII: 4659. Additionalcollection: KBA:"Sambas River", 30. 7. 1903, Micholitzs.n. (H-BR).

Leucophanescandidum (Schwaegr.) Lindb. in Ofvers.F6rh. Kongl. Svenska Vetensk.-Akad. 21:602 (1865). -Syrrhopodon candidusSchwaegr., Sp. Musc.Frond. Suppl. 2: 105 (1827).-Typus: Indonesia,Java?"Ex insula Iavania?",Reinwardt s.n. (Lecto-G! isolecto-BM, G!).

Ic.: Schwaegrichen1811-1842: pl. 182, figs. 1-8 (as Syrrhopodoncandidus); Dozy & Molkenboer1854- 1870:pl. 16 (asL. reinwardtianum);Cardot 1900: pl. 8,figs.43a,b,pl.9, figs.43c-q (asL.fusculum), fig. 45a (as L. densifolium),pl. 10,figs. 45b, c (asL. densifolium);Fleischer 1904: 182,fig. 24; Brotherus1924: 224, fig. 179; Bartram1939: pl. 5, fig. 79; Johnson 1964: 356, figs. 13a, b, fig. 13f (as L. naumanni).

Plantsup to 6(-7) cm high,glossy, growing in densetufts or cushions.Leaves up to about4 mm long, base sheathing,obovate, upper parts broadly lanceolate, keeled, mostly spreadingor squarrose,some- times erect.Leaf margins entire below, toothed above,apices mostly sharplyapiculate. Hyaline lamina short, 1-3 cells wide nearleaf base.Costal hyalocystsin one layeradaxially and abaxiallythroughout leaves. Dioicous. Femalegametoecia terminal (pseudolateral). Seta up to 1.3 cm long, smooth. Capsule about 1 mm long, erect,cylindrical. Male plants not observed. L. candidumis readilyrecognized by the mostly squarroseleaves with sheathingbases and usually sharplyapiculate apices. The leaf structureis similarto thatof L. octoblepharioidesin that the costalhyalo- cysts are in one layeron eitherside of the chlorocystlayer throughout the leaves. This speciespreferably grows epiphytically, but also on decayingwood, soil, and differentkinds of rocks.It is quite common in lowlandforests up to about 1000 m, but rapidlybecomes rare above that al- titude.

Geographicalrange (from SalazarAllen 1986): Malaysia,Indonesia (Borneo: SAB, SAR,KBA, KTI, KSE),Philippines, New Guinea,Solomon Islands,Australia (Queensland), islands of the tropicalPacific. Altitudinalrange (Borneo; based on literature):approximately 100-1150 m. Specimensexamined: BRYOTROP:none. Additionalcollections: SAR: Lundu,23. 8. 1903,Micholitzs.n. (H-BR).-KBA: Lebang,Hara,130 m,2. 1. 1925, Winklers.n. (H-BR). - KSE:Bandjarmassin, Tamnalagang, com. Grebe s.n. (H-BR). Wilidenowia18 - 1989 547

Leucophanesglaucum (Schwaegr.) Mitt. in J. Proc.Linn. Soc. Bot. Suppl. 1: 25 (1859). -Syrrhopodon glaucusSchwaegr., Sp. Musc. Frond.Suppl. 2: 103 (1827). Typus:Marianas, Gaudichaud 10 (Lecto-G!, isolecto- BM, G, L); Indonesia, Moluccas, "in Insula Rauwack Moluccarum...",Gaudichaud s.n. (Syn-G). = Leucophanesalbescens C.Muell. in Bot.Zeitung(Berlin) 22:347(1864).-Typus: Malaysia,"Pulo Penang IndiaeOrientalis: Didrichsen Exp. transatlantica Danica",J. Langes.n. (Iso-BM,FH,S) -cf. SalazarAllen 1986. = Leucophanesglaucescens C. Muell.ex Fleisch.,Musci Buitenzorg 1: 178(1904). - Typus:Indonesia,Java, "ImBot. Gartenzu Buitenzorgnicht selten,aber meist steril",Fleischer s.n.; Sumatra,"bei Palembang am Lemattang,100 mi, Fleischers.n.;"FernerOst. Indien (ex Herb. Brit. Mus.) c. fr.",Kurz(all specimens FH) - cf. SalazarAllen 1986. = Leucophanessubglaucescens C. Muell. ex Dix. in J. Linn.Soc. Bot. 43: 296 (1916).-Typus:Indonesia, Ti- mor, Micholitz8 (Lecto-BM! isolecto-BM, FH, H-BR!, S) - cf. SalazarAllen 1986. - Leucophanescompactum Broth. in C. Muell.,Genera Musc. Frond. 86 (1900),nom. nud. - Originalcol- lection: Malaya,Great Natunas, 1. 8. 1892, Micholitz82 (duplicateH-BR!). = LeucophanespugionatumC. Muell. in GeneraMusc. Frond. 86 (1900),nom. nud.- Originalcollection: Indonesia,Timor, "Timorlaut, ins. Sarak",20. 4. 1891, Micholitz7 (duplicateH-BR!). = Leucophanessubalbescens Broth. in Nat. Pflanzenfam.1(3): 347 (1901), nom. nud.- Originalcollection: NicobarIslands, Kurz 3853, 3854 (duplicatesin H-BR!).

Ic.:Schwaegrichen 1811-1842: pl. 181(as Syrrhopodonglaucus); Geheeb 1898: pl. 5 (asL. beccarii);Cardot 1900:pl. 10, figs.47a-c; Miller,Whittier&Bonner 1963: pl. 5 (asL.glauculum); Miller& Smith 1968:230, figs.5a-d,231, figs.7a-c (asL.glauculum); Gangulee 1971: 433, fig.203 (asL.glaucescens), 435, fig.204 (as L. albescens);Whittier 1976: 99, figs. 25D-F (as L. albescens).

Plantsup to 3 cm high,growing in ratherdense to verydense tufts or cushions.Leaves up to 4.5 mm long, spreadingto curvedor nearlyerect, lanceolate from oblanceolate, semisheathing base, upperparts keeledto channelled,often twistedwhen dry.Leaf margins entire below, becoming serrulate and gradual- ly more stronglyserrate upwards, apices mostly acute, apiculate, toothed, apical gemmae common. Hya- line laminavanishing at or belowmidleaf. Costal hyalocysts near leaf base in 1 layeradaxially and in 2-3 layersabaxially, above in 1 layeron eitherside. Dioicous.Female gametoecia terminal (pseudolateral). Seta up to 7 mm long, smooth.Capsule 1.2- 1.3 mm long, erect, cylindricalor narrowlyellipsoid. Male plants not observed. Once familiar,L. glaucumis rathereasily distinguished by certainmore or less difficult-to-defineha- bitualcharacters. However, the structureof the leavesis uniqueamong the BorneanLeucophanes species: the abaxialcostal hyalocystsnear the leaf bases are in 2-3 layers.Hence, the speciescan positivelybe identifiedby cross-sectionsthrough the basal portions of the leaves. SalazarAllen (1986) recognized3 subspecieswithin the variableL. glaucum,but since only subsp. glaucumis reportedfrom Borneo, I will not makefurther comments on thesein the ;presentpaper. Among the mostvariable characters in L.glaucum are the leafshape, the abaxialspiculosity of the centralstereids, and the peristomeornamentation. The lastmentioned character appears to have some taxonomicvalue at subspecificlevel. Unfortunately,no altitudinaldata for L. glaucumare given in the literature.However, the speciesis obviouslyrestricted to coastalareas and verylow latitudes.In New Guinea,for example,L. glaucumhas been collectedfrom sea level up to just20 meters.It most commonlygrows on treetrunks and bases,but occasionallyalso on decayingwood, soil, and rocks. I had accessto only one Borneanspecimen of L.glaucum, and the abovedescription is mainlybased on New Guineanmaterial. 548 Enroth:The bryophytesof Sabah(North Borneo)IV.

Geographicalrange (from Salazar Allen 1986):"... from Indiato Malesia,Japan,northeastern Australia, Vanuatu,New Caledonia,the islandsof the tropicalPacific and a disjunctpopulation in Guadeloupe (WestIndies)". Borneo: SAB, SAR,KSE. Altitudinalrange: badly known, but obviouslyrestricted to very low altitudes. Specimensexamined: BRYOTROP:none. Additionalcollections: KBA: "Prope Pontianak", Oorschot s.n. (H-BR).

Leucophanesoctoblepharioides Brid., Bryol. Univ. 1: 763 (1826). Typus:Indonesia, Java (Lecto-B!, isolecto-BM); Nepal (Syn-B!). = Leucophaneskorthalsii Dozy &Molk., Musci Frond. Ined. Archip. Ind. 3: 65 (1846) Leucophanesoctoble- pharioidesvar. korthalsii (Dozy&Molk.) Fleisch.,Musci Buitenzorg 1: 176(1904). -Typus: Indonesia,Bor- neo, KalimantanSelatan, "prope pagum Martapoera", Korthals s.n. (Lecto-L); Indonesia, Amboina, Zip- peliuss.n. (Syn- L).- cf. SalazarAllen 1986. - Octoblepharumsmaragdinum Mitt. in Bonplandia9: 366 (1861) Leucophanessmaragdinum (Mitt.)Jaeg. in Ber.Thitigk. St. GallischenNaturwiss. Ges. 1877-1878:391 (1880).- Typus:Fiji, Seeman 863 (Holo- NY!).

Ic.: Schwaegrichen1811-1842: pl.311 (asSyrrhopodon octoblepharis); Dozy&Molkenboer 1845-1848: pl. 23 (asL. korthalsit);Cardot 1899: pl. 1,fig. 1; Cardot1900: pl. 11,figs. 52a-e; Bartram1939: pl.6, fig.81; Johnson 1964:356, fig. 13d,357,figs.14a-c; Gangulee 1971: 436,fig.205,438,fig.206 (as L. nicobaricum); Whittier1976: 99, figs. 25G-I.

Plantsup to about4 cm high,growing in loose to densetufts or cushions.Leaves up to 6.5 mm long, erectto patent,lanceolate from oblanceolatebase, sometimes nearly linear, keeled below, apices obtuse to acute,usually dentate, apical gemmae common on leaves.Costal hyalocysts adaxially and abaxiallyin one layerthroughout the leaves. Dioicous. Femalegametoecia terminal (pseudolateral). Seta up to 11.5 mm long, smooth.Capsule 1.3-1.5 mm long, erect,cylindrical to narrowlyelliptical. Male plants not observed. L. octoblepharioidesis quite a common speciesin Indonesiaand adjacentregions. It has the costal hyalocystsin one layeron eitherside of the chlorocystlayer throughout the leaves,whichserves to distin- guishit fromL.glaucum and L. angustifolium.L. candidum,whichhas a similarleaf structure,is moreglos- sy and has more stronglykeeled leaves, narrowing from a sheathingbase into a mostlysquarrose upper part. Forsome reasonSalazar Allen (1986) failed to locate the type specimenof L. smaragdinum,but I managedto receiveit on loan fromNY. L. smaragdinumis nothing morebut a typicalform of L. octoble- pharioides;the formername has been misapplied,particularly in Oceania,for examplein Miller&Smith (1968),for plants that obviouslybelong to L. glaucum. Only one Borneanspecimen studied by the presentauthor bore sporophytes. The capsules were only 1.3-1.5 mm long, whereasSalazar Allen (1986)gives a length rangeof (1.5)1.7-2.6mm for this species. L. octoblepharioidesis most common in lowlandforests below about 1000 m. It growsepiphytically on tree trunksand tree bases,but also on decayingwood, rocks,and soil.

Geographicalrange (from Salazar Allen 1986):Seychelles, Mauritius, Sri Lanka, Nicobar Islands, Nepal, SouthernJapan, Taiwan, Vietnam, Thailand, widely distributed from Malaysia through Indonesia (Bor- neo: SAB,SAR, KBA, KSE), New Guinea,Philippines, Australia (Queensland), New Caledoniaand the tropicalPacific. Willdenowia18 - 1989 549

Altitudinalrange (Borneo): up to 900 m. Specimensexamined: BRYOTROP: V: 4472. - IX: 4366. Additionalcollections: SAR: s. loc., nativecollector 1200 (H-BR).- KBA:"Prope Pontianak", Oorschots.n. (H-SOL). - KSE:Bandjarmassin, Tamnalagang, com. Grebes.n. (H-BR).

ArthrocormusDozy & Molk., Musci Frond.Ined. Archip.Ind. 75 (1846).

Thisgenus comprises only one species,A. schimperi(Dozy& Molk.) Dozy&Molk., distributed from Sri Lankaand Thailandto the SocietyIslands in the Pacific.

Arthrocormusschimperi (Dozy & Molk.) Dozy & Molk., Musci Frond.Ined. Archip. Ind. 76 (1846). --Mielichhoferia schimperiDozy & Molk. in Ann. Sci. Nat. Bot., ser.3, 2: 312 (1844). Typus:Indonesia, Borneo, Kalimantan Selatan, "prope Martapoera", Korthals s.n. (Holo- L, iso- H-BR!, H-SOL!).

Ic.: Dozy& Molkenboer1845-1848: pl. 27; Cardot1899: pl. 1, fig. 12 (asA. thraustus);Cardot 1900: pl. 14, figs. 68a-d, pl. 15, figs. 69a-h (as A. thraustus);Fleischer 1904: 185,fig. 25; Brotherus1924: 227, fig. 181; Bartram1939: pl. 6, fig. 83;Johnson 1964: 359, fig. 16; Miller&Smith 1968:231, fig. 8; Onraedt 1976:365, fig. 10: 2; Whittier1976: 110, fig. 28.

Plantsmostly 1.5-3.5 cm high, growingin ratherloose tufts or individuallyamong other mosses. Leavesup to 8.0(-9.0) mm long, in 3 distinctranks, subulate from oblanceolate, keeled based, thick and stiff but fragile with the tips mostly brokenoff, leaf apices acute, apiculate;gemmae not common. Hyalinelamina vanishing below 1/2 leaflength.Costal chlorocysts in one layernearleafbases, in 2 layers upwardsand finallyin 3 layersabove, superficial chlorocysts embedded under a hyalocystlayer of one cell in thickness. Dioicous.Female gametoecia terminal. Seta 7-8 mm long, smooth.Capsule 1.0 mm long, erect, cy- lindrical.Male plants not observed. A. schimperiis very distinct in its 3-ranked,stiff but fragileleaves with the tips usuallybroken off. Gemmaeon the leavesare not frequentlyproduced and theywere reported for this speciesonly recently (Enroth 1988). A. schimperiis restrictedto low altitudes,where it is quite common in suitablehabitats. A quotation fromMeijer (1954) may be pertinent:"On the drierhills betweenthe rivuletsepiphytes and epiphyllous speciesare rather scarce. Looking at the bryofloraI could tell whetherwewere near a riveror not. On the drierhills almost the only substrataworth searching for bryophytesare the trees,many of them belong- ing to the Dipterocarpaceae,but severalother families are also present. The most common specieson the rathershady bases of these is Arthrocormusschimperi Dz. & Mb.".

Geographicalrange: Sri Lanka,Thailand, widely distributed in Malaysiaand Indonesia(Borneo: SAB, SAR,KBA, KTI, KSE), Philippines, New Guinea,Solomon Islands,Australia (Queensland), Carolines, Vanuatu,Fiji, Samoa, Society Islands. Altitudinalrange (Borneo): 20-500 m. Specimens examined: BRYOTROP:Ia: 4555, 4582. Additionalcollections: SAR: Gunong Mulu Nat. Park,alluvial forest NE of BaseCamp, along trailto LongBerar, basal part of heavytree trunk, full shade,moist, 2.5. 1978,Touw 19498 (H,L). - KBA:Sambas, Siluas,22.7.1903, Micholitzs.n. (H-BR).- KSE:Bandjarmassin,Tamlanagang, com. Grebes.n. (H-BR). 550 Enroth:The bryophytesof Sabah(North Borneo) IV.

ExostratumL. T. Ellis in Lindbergia11: 22 (1985).

In his revisionof the genusExodictyon Ellis (1985) segregated Exostratum from it, mainlydue to the differentarrangement of the superficialcostal chlorocysts. In the formergenus these form a networkbe- tweenthe costalhyalocysts,whereas in the latterthey are arranged to forma continuouslayer (this differ- ence is diagrammaticallyillustrated on page 18 in Ellis'paper). I agreewith Ellis and think that Exodictyon andExostratum have to be segregatedat genericlevel, although Cardot (1900) considered these taxa only as sections. Exostratumcomprises four species,two of which have been reportedfrom Borneo.

Key to the Borneanspecies of Exostratum

1. Projectionsof costa in subulaup to about 15 pm high ...... E. blumei - Projectionsof costa in subulaup to 30-35(-40) pm high ...... E. sullivantii

Exostratumblumei (Nees ex Hampe) L. T. Ellis in Lindbergia11: 25 (1985). 8 Syrrhopodonblumii Nees ex Hampe in Bot. Zeitung(Berlin) 5: 921 (1847). Typus:Indonesia, Java, Blume s.n. (Lecto-L, isolecto-BM!, NY).

Ic.: Dozy& Molkenboer1854-1870: pl. 16 (asLeucophanes blumii ("Blumii')); Cardot 1899: pl. 1, fig. 14 (asExodictyon nadeaudii ("Nadeaudi')); Cardot 1900: pl. 17,figs. 74a-e (asExodictyon scabrum),figs. 75a-d (asExodictyon arthrocormoides), figs. 77 a-c (asExodictyon nadeaudii ("Nadeaudi")); Fleischer 1904: 189, fig. 26 (asExodicyon blumii); Brotherus 1924: 227, fig. 182B(as Exodicyon nadeaudii ("Nadeaudii')); Bartram 1939:pl. 6, fig. 85 (asExodictyon blumii ("Blumii"));Johnson 1964: 360, fig. 18 (asExodictyon blumii); Gan- gulee 1971: 445, fig. 208 (as Exodictyonblumei); Ellis 1985: 26, fig. 10, 27, fig. 11, 28, fig. 12. Map: Ellis 1985: 29, fig. 13.

Plantsmostly 1-3.5 cm high, growingin loose to rathercompact tufts or mats.Leaves in 3 obscure ranks,mostly 3-4.5(-5.5) mm long, erectopatentto patent,subulate from an oblanceolate,semisheath- ing base, apices obtuse to acute, toothed; apical gemmae common. Leaf marginsentire at base, at shouldersusually ciliate, more or less stronglytoothed in the subula.Superficial costal chlorocystsex- posed, often drawnout into single spinose projectionsup to about 15 pm high, usuallylower. Dioicous. Gametoeciaterminal. Sporophytes not observed. E. blumeihas clearlylower projections of the costal chlorocystsin the subulathan E. sullivantii(see the keyabove). Arthrocormus schimperi, on the otherhand, has the leavesdistinctly 3-ranked with the tips mostly brokenoff, and a smooth surfaceof the subula. E. blumeigrows on a varietyof substrates:tree trunksand roots, decayingwood, soil, and cliffs or rocks.It favorsshady habitats, preferably with high humidity,and is distributedin lowlandand lower montane forestsup to about 1700 meters.

Geographicalrange (from Ellis 1985):India, Sri Lanka, N Vietnam,Thailand,Malay Peninsula, Sumatra, Java,Borneo (SAB, SAR,KBA), Sulawesi, Moluccas, Taiwan, Batan Islands, Philippines, New Guinea, Australia(Queensland), Fiji, Samoa, Society Islands. Altitudinalrange (Borneo): 150-1700 m. Specimensexamined: BRYOTROP:XII: 4688. - XVI: 4254. - XVIa:3601. Additionalcollections: KBA: Sungei Malang,28. 1. 1924, Winklers.n. (H-BR). Willdenowia18 - 1989 551

Exostratumsullivantii (Dozy & Molk.) L. T. Ellis in Lindbergia11: 29 (1985). - Syrrhopodonsullivantii Dozy & Molk., Bryol.Jav. 1: 58 (1856). Typus:Indonesia, Java, Holle s.n. (Holo- L, iso- BM!, L).

Ic.: Dozy& Molkenboer1854-1870: pl. 47 (asSyrrhopodon sullivantii ("Sullivanti'"); Cardot 1900: pl. 16, figs.73a-k (asExodictyon hispidulum); Bartram 1939: pl. 6,fig. 84 (asExodictyon sullivantii);Johnson 1964: 359, fig. 17 (as Exodictyonsullivantii); Ellis 1985: 30, fig. 14, 31, fig. 15. Map: Ellis 1985: 29, fig. 13.

Plantsmostly less than 1.5(-2.0) cm high, formingrather loose tufts or mats.Leaves in 3 obscure ranks,up to 3.0(-3.2) mm long, hispid,subulate from oblanceolatebase, subulae mostly spreading and recurved,apices obtuse to acute,strongly toothed; apical gemmae common. Leaf margins entire at base, usuallyciliate at shoulders,strongly toothed with spinose teeth above. Chlorocystsof the subulain 3 layers,those of the costa surfacemostly drawn out into spinose,up to 30-35(-40) pm high projections. Dioicous. Femalegametoecia terminal. Sporophytes and male plants not observed(description based on New Guinean(kept in H). E. sullivantiiwas not includedin Touw's(1978) list ofBorneanmosses, and, as far as I know,it wasfirst recordedfor Borneo by Ellis(1985). I havenot seen the specimencited by him,i.e. EastKutai, peak ofBa- likpapan,terrace Beul, Meier B2067 (L),but thereis no reasonto doubt the correctnessof his identifi- cation. Accordingto Ellis (1985) E. sullivantiigrows on the trunksof trees and tree ferns,and decaying stumps.He givesthe altitudinalrange of 1400-2900 m, but on the Huon Peninsula,Papua New Guinea, the specieswas collected between 900-1300 m. Hence, E. sullivantiiobviously occurs mainly in lower and uppermontane forests.

Geographicalrange (from Ellis 1985):Java, Borneo (KSE),Sulawesi, New Guinea,Philippines. Altitudinalrange (based on literatureand specimensfrom New Guinea):900-2900 m. Specimensexamined: BRYOTROP:none.

OctoblepharumHedw., Sp. Musc. Frond.50 (1801).

Octoblepharumis a slightlyanomalous genus within the Calymperaceaesubfamily Leucophanoideae, re- gardingboth the gametophyteand the sporophyte.The chlorocystsin the subulaare triangular in cross section and arrangedin a single layer,and the leaves lack a marginalrib. In addition,the peristome deviatesquite strongly from otherLeucophanoideae: the numberof peristometeeth is only8, andtheyare low,broad, and nearlysmooth on the surface.However, 0. albidumhas been reportedto producetypical "calymperoid"gemmae (Harrington & Egunyomi 1976). The leaf structureof 0. albidumwas studiedin detail by Ligrone& Egunyomi(1981-1982). In the palaeotropicsOctoblepharum is obviously represented by only one species,0. albidumHedw., but in SouthAmerica the numberof speciesis probablyhigher. For example, Schultze-Motel & Menzel (1987) recognize5 species for Peru.

Octoblepharumalbidum Hedw., Sp. Musc. Frond.50 (1801). Typus:Jamaica, Swartz s.n. (Holo- G!). = Octoblepharumlongifolium Lindb. in Ofvers.Forh. Kongl. Svenska Vetensk.-Akad. 21: 608 (1865).-Ty- pus: Society Islands("Tahiti"), Ponten s.n. (Holo- H-SOL!). 552 Enroth:The bryophytesof Sabah(North Borneo)IV.

Ic.: Cardot1899: pl. 1,fig. 10;Cardot 1900: pls. 12-14, figs.61a-r; Fleischer 1904: 170,fig. 22; Brotherus 1924: 226, fig. 180, 227,fig. 182A;Bartram 1933: 58, fig. 37; Bartram1939: pl. 6, fig. 82; Schornherst Breen1963: pl. 24, figs.2,3,5-8;Johnson 1964:358, fig. 15;Gangulee 1971: 441,fig. 207;Onraedt 1976: 365, fig. 10: 3; Whittier1976: 106, figs. 26A-C, figs. 26D-F (as 0. longzfolium).

Plantsless than 2 cm high, growingin ratherdense to loose cushions or mats. Leavesup to 9(- 10) mm long, often shorter,thick, usually more or less recurved,rarely erect, narrowly ligulate from ob- lanceolatebase, apices rounded to obtuse,apiculate, often brownish.Leaf margins entire throughout or serrulateto serratenear apices. Hyaline lamina confined to 1/1-1/5 of the leaf lengthfrom base. Costal hyalocystsin 2-5 layerson eitherside of the chlorocystlayer in upperparts of leaves.Costal chlorocysts mostly triangularin cross section, arrangedin a zig-zagpattern. Autoicous, fruitingcommon. Gametoecialateral. Seta 3.5-5.0 mm long, smooth. Capsule 1.0(- 1.2) mm long, erect, cylindrical. 0. albidumis a peculiarplant, and may be easilyrecognized by the habit alone, but since habitual charactersare often moreor less difficultto defineand describe,I preferto stresssome microscopicchar- actersof diagnosticvalue. The hyalinelamina lacks a marginalrib; the costalhyalocysts are in 2-5 layers on eitherside of the chlorocystlayer in the upperparts of leaves;the costalchlorocysts are triangular and form a typical zig-zagpattern as seen in cross section. 0. albidumis a lowlandspecies, rarely occurring above 1500 m and obviouslymost common below approximately1000 m. Treebases and lowertrunks in shadyplaces are the preferredsubstrates, but the species has also been collected on decayingwood.

Geographicalrange: pantropical at low altitudes.Borneo: SAB, SAR,KBA, KSE. Altitudinalrange (Borneo): c. 150-1300 m. Specimensexamined: BRYOTROP:XIII: 3027, 3055. Additionalcollections: SAR: Gunong Mulu Nat. Park,temporarily dry branch of MelinauR. near Camp 5 (MelinauGorge), low on treetrunk, full shade,moist, 17.5.1978,Touw20572 (H, L).-KBA"PropePon- tianak",Oorschot s.n. (H-BR). - KSE: Bandjarmassin,com. Grebe s.n. (H-BR).

Acknowledgements

I am gratefulto Prof.Dr. Wolfram Schultze-Motel and Mr. Mario Menzel (Botanisches Museum und BotanischerGarten Berlin-Dahlem), who offeredme the opportunityto contributeto the BRYOTROP project.The Curatorof BM is acknowledgedfor sending essentialspecimens on loan.

References

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Addressof the author: JohannesEnroth, Botanical Museum, University of Helsinki,Unioninkatu 44, SF-00170 Helsinki, Fin- land.