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Bull. nat.Hist. Mus.Land. (Bot.)30(2): 33-79 Issued30November2000 The genus Polystichum (Dryopteridaceae) in " Africa ./ . \") , JACOBUS P. ROUX 22 NOV NationalBotanicalInstitute, ComptonHerbarium, PrivateBagX7, Claremont7735, SouthAfrica igfgKAL LJ6RARV CONTENTS Introduction 33 Materialsandmethods 34 Terminology 34 Taxonomictreatment 35 KeytotheAfricanspeciesofPolystichum(includingMarionandPrinceEdwardIslands) 36 References 78 Systematicindex 79 SYNOPSIS. PolystichumRothisaferngenusof160to200speciesoccurringthroughoutthetemperatepartsoftheworldandthe montanetropics,butabsentfromthearidregionsaswellasthelowlandtropics.AlthoughfloristicaccountsofPolystichumexist formanypartsoftheworld,thegenusremainspoorlyunderstoodandataxonomicaccountforthegroupasawholehasnever beenundertaken.PolystichumispoorlyrepresentedinAfrica(includingtheMarionIslandgroup)withmerely16speciesandone knownhybridhavingbeenrecordedfromtheregion.Mostofthespeciesareconfinedtosub-SaharanAfrica,occurringmainly alongtheeasternmountainrangesandMt. Camerooninthewest. InthisreviewoftheAfricanPolystichumspeciesdetailed observationsarepresentedespeciallyonthepaleae, asthese structureswerefoundtoprovidethebestcharactersonwhicha subgenericclassificationcanbebased.Diagnosticfeaturesandrelationships,variation,andthedistributionandecologyofeach speciesareincluded. INTRODUCTION allopolyploids (Vida& Reichstein, 1975; Wagner, 1979). Alsothe frequentoccurrenceofFl-hybridsinsomegroupsobscuresspecies limits. The genus Polystichum Roth consists of between 160 (Tryon & Aformalsubgenericclassificationforthegenushasthusfaronly Tryon, 1982)and200(Daigobo, 1972)species.Itoccursthroughout been provided for the east Asian species by Tagawa (1940) and thetemperatepartsoftheworldaswellasthemontanetropics,but Daigobo(1972).Someofthesesectionshavesincebeensubdivided is mostly absent from the lowland tropics. Within the range two further by Zhang & Kung (1995, 1996a, b) and Kung & Zhang distinctcentresofdiversitycanbeidentified,namelyalargerAsiatic (1998) to make provision for some ofthe Chinese species. Since centrewithapproximately70speciesandatropicalAmericancentre mostspeciesremainpoorlyknownaphylogenyforthegenuscannot with approximately 55 species. Both these regions fall within the beproposed. mountainoustropicscharacterizedbymild,moistclimatesthatare Althoughfloristicaccountsofthegenushavebeenpublishedfor hardly seasonal. They also correspond with the areas of highest manypartsoftheworld,somebeingveryoldandoutdated,nosingle speciesdiversityforhomosporousferns (Tryon, 1985). monographictreatmentexistsandmostspeciesremainpoorlyknown. WithintheDryopterideaePolystichumismostcloselyrelatedto Some ofthe modern-day regional treatments provide no detailed Arachniodes Blume, Cyrtomium C. Presl, Dryopteris Adans. and observations that may suggest affinities. Within the study area Phanerophlebia C. Presl. Cyrtomium and Phanerophlebia have floristic accounts for Polystichum are available for North Africa often been included in Polystichum (Kramer, 1990), but it has (Maire, 1952), West tropical Africa (Alston, 1959), Cameroon subsequently been shown that both these genera have a closer (Tardieu-Blot, 1964), Mozambique, Malawi, Zambia and Zimba- affinitytoPolystichumthantoeachother(Yatskievych, 1996). bwe (Schelpe, 1970), Rwanda, Burundi and Kivu (Democratic Polystichuminastrictsenseisanaturalandrelativelyhomogene- RepublicofCongo)(PichiSermolli,1985),southernAfrica(Schelpe ousgroupofplantscharacterizedby 1-pinnateto3-pinnatelaminae &Anthony, 1986) andBioko(Benl, 1991). withacroscopicallydevelopedultimatesegments,anadromousfree It is a well-known fact that Africa, when compared with other venation, uni- orbiseriatecircularsori positioned medially, termi- tropical parts of the world, supports a floristically impoverished nally or near terminally on abbreviated or unabbreviated vein vascularflora.Thisphenomenonisalsoreflectedinthepteridophyte branches,andpeltateindusia(anumberofspeciesareexindusiate). floraofthecontinent.Thecauseofthisfloristicpovertyisascribed Thereceptacleappearstobenudeinmostspecies. to the isolation ofAfricafrom the othercontinents since the mid- Although Polystichum as a genus is easily recognized, many Cretaceousandthesubsequentsignificantchangesintheclimateas speciesaresuperficiallyverysimilar.Thedelimitationofspeciesis aresultofuplift,continentaldriftandaridificationcausedbyextra- furtherhamperedbythefactthatmany,mostlycommonspeciesare tropical glaciation. All these changes may well have resulted in a TheNaturalHistoryMuseum,2000 34 J.P.ROUX progressive elimination of the once rich tropical and subtropical mountain systems, whilstP. transvaalense and P. wilsonii are dis- foreststhatexistedtowardsthelateJurassicandtheestablishmentof tributedthroughoutseven mountainsystems.ThedistributionofP. extensivedeserts and semi-desertsbytheearly Pliocene (Coetzee, wilsonii, however, also extends along the Himalaya mountains to 1993). Bhutan, Japan and Taiwan. Polystichum luctuosum has an almost Within the study area two regions can be identified: an African similareastern distribution butis confinedtotheDrakensberg and and a sub-Antarctic region. The origin, composition and floristic Chimanimani mountain systems in Africa. Polystichum luctuosum affinitiesoftheseregionsdiffermarkedly.TheAfricanregionisthe andP.wilsoniialsoshowadisjunctdistributionwiththeMadagascan largest and today there are three fundamentally different floras or region. biogeographical subregions which can be identified: a southern Basedonobservationstaken fromthesystematic treatment, and Africanflora,atropical AfricanfloraandaNorthAfricanflora. judging from wide-ranging species, southern and tropical African The flora ofthe southern African subregion is believed to have PolystichumhasacloseraffinitywithtaxafromAsiathanwiththose evolved gradually since the mid-Tertiary, derived partly from an from the Americas. Polystichum in the North African phyto- ancientsouthernAfricantemperatefloraandpartlyfromatropical geographical subregion shows a closer affinity with Polystichum African forest flora (Goldblatt, 1978). Elements of the southern fromEuropethanfromAfricaasP. aculeatumandP. setiferumare AfricanfloracurrentlyextendintotropicalAfricaalongtheeastern widespreadinthatregion.OnlyP. marionense,endemictoMarion, escarpment.ThesouthernCapeforestsarebelievedtobeimpover- Prince Edward and Crozet Islands, occurs in the sub-Antarctic & ished remnants of the tropical African forest flora (Coetzee phytogeographical region. Muller, 1984). Also the tropical African flora is believed to be an impoverishedremnantofaoncemuchrichertropicalrainforestflora thatextendedoverafargreaterareathanitcurrentlyoccupies. MATERIALSANDMETHODS North Africa has also experienced significant changes in its climate and vegetation. During the Palaeocene the present Sahara desert was clothed by a rich tropical lowland rainforest that also This review is basedon observations made duringextensive field- coveredpartofEurope(Greenway, 1973;Raven&Axelrod, 1974), work in southern Africa and on cultivated plants collected during butbytheOligo-Mioceneitwasreplacedbyasubtropicalwoodland thesetravels.Thecollectionsofseveralherbariawerealsostudied. cslaivmaantneaes(tAaxbellirsohded&itsRelafveinn,the19m7a8)j.orFpraortmotfhethePlSiaohcaernae (aQudeezseelr,t TNhHe,seNUin,clPu,dPe:REB,,BRMA,B,BOSLA,M,BRS,REGTHH,,WGRAAG,(Ka,bLb,reMv,iaMtiAoLns,fNoBllGo,w 1978), serving as an effective barrierto migration from the south. Holmgren et al., 1990) and the private herbarium ofProf. R.E.G. The formation of glaciers on the high mountains during the Pichi Sermolli (PIC.SERM.). Pleistocene permittedtheestablishmentofcircumborealelements. Paleaandindusiumobservationsweremadebyremovingasmall ManyoftheseelementsarepresentintheNorthAfricanflorasince number of these structures from selected specimens. These were it is composed of relict elements of African origin as well as cleaned and cleared in diluted household bleach, after which they elements from Eurasia, not frequent in the present sub-Saharan were semi-permanently mounted in glycerine and the cover slips flora. The mediterranean influence on the flora of North Africa sealed with Entellan. Observations were made with an Olympus justifies it being considered as a biogeographical subregion ofits CH-2light-microscopefittedwithadrawingtube. own. Thecollections studiedareall listedunder 'Materialexamined'. The Marion and Prince Edward Island group forms part ofthe These are arranged alphabetically according to country oforigin. sub-Antarcticregion,aphytogeographicalareacompletelydifferent South African (including Lesotho and Swaziland) collections are from the foregoing. This island group is ofvolcanic origin and is furtherarr&angedaccordingtothequarter-degreesquare-gridsystem estimatedtobe0.5 millionyearsold(Verwoerd, 1971). Situatedin (Edwards Leistner, 1971).Inthissystemeachone-degreesquare theSouthernOceansome 1800kmfromAfrica,itsbiotaconsistsof is known by a standardized name, derived from a town or other ttahxeAambscoeaulptvaebtslwoei-nothfhailbroidtnsagot-sfditnshotetaAnafclrewiadcyiassnpfepratsveaorluirdaaonbpdlheytthfeeosraabprilelainlttiymgitrtooewedstithna.btlhiesihr dfaeinavdtiuDdreedfroionftmoilmfepoftuorrttohaarnlicfgeh-tdeiagnnrdetehteospqsutqaouraberoset.(t3oE0m'a.cxEh3a0co')hn,eh-nadulemfg-brdeeeregerdeseqAus,aqrBue,arCies ohcocwuervreern,cearteoctlhoesecloyntrielnaetnetd(tKoortanxaas,in1e9i9t3h)e.rTthroepimcaajloArimteyriocfathaensde/, nisumagbaeirnedsuAb,diBv,iCdeadndinDt.oBqyuaurstienrg-dtehgerseeecos-qouradriensat(e1s5'axgeo1g5'r)a,phaigcaailn orsoutheastAsia.Pteridophytesofthesub-Saharabiogeographical areacanimmediatelybeidentified. regionexhibitthreediscontinuousdistributionpatterns: an Ameri- Unlesscitedotherwise, thechromosomenumbersprovidedhere can-Africandisjunction,anAfrican-Madagascandisjunctionandan are based on the author's own observations and will be published African-Asiandisjunction. elsewhere. Polystichum in Africa is largely confined to the Afromontane Phytochorion. White (1978)dividedthismontanearchipelagointo Terminology sevenregional mountain systems. TheNorth African Atlas moun- This study is principallybasedon adetailedcomparative morpho- tain ranges arehereaddedasaneighth. Althoughthe sub-Saharan logicalanalysisofthe sporophyte, wherepaleastructureprovedto mountainrangesaresufficientlydistinct,thesystemsareconnected bemostinformativeinsuggestingspeciesgroups.Thetermsusedto byacomplexseriesofintermediatefloras(Fig. 1).TheDrakensberg describetheapexortheapicalcellofthepaleaearedefinedas: system, with six Polystichumendemics, is the richest. This is also true for the angiosperms (White, 1978). The only other mountain - apexflagelliform:theapexofthepaleaterminatesinauniseriate systemswithtrueendemicsaretheImatongs-Usambarasystemwith seriesofslendercells. twoendemics(P. kilimanjaricum&P. volkensii)andtheEthiopian - subulatecell:theapicalcellislessthan0.4mmlongandtheapex systemwithoneendemic(P.magnificum).OtherAfricanPolystichum isusuallyblunt. specieshavewiderdistributions.Polystichumzambesiacumoccurs - acicularcell:theapicalcellisslender,straight,morethan0.4mm in the Chimanimani, Uluguru-Mulanje, and Imatongs-Usambara long, andtheapex isusually sharp. POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 35 V r VIII. NORTHAFRICAN 1 IV. IMATONGS-USAMBARA U-y\P.kilimanjaricum VII. ORAKENSBERG '.drakensbergense .incongruum macleae . .monticola .pungens .transkeiense Fig. 1 DistributionofPolystichuminAfrica. filiformcell:theapicalcellisslender,twisted,andmorethan0.6 HemesteumH.Lev.,Fl.Kouy-Tcheou:450,496(1915),nonNewm. mm long. (1851). Type species: severalPolystichumspeciesarelisted. thin-walledcell:thesecellscanvaryconsiderablyinsizeandthe Aetopteron HouseinAmer. FernJ. 10: 88 (1920), nom. nud. cell wall is conspicuously thinner than that ofthe surrounding Papuapteris C. Chr. in Brittonia 2: 300 (1937). Type species: cells, and is therefore often lost. In dry material the cellular Papuapteris linearis C. Chr. (= Polystichum lineare (C. Chr.) contents areusuallyyellowishincolourandappearcrystalline. Copel.). Acropelta Nakai in Bull. Natl. Sci. Mus. Tokyo33: 5 (1953). Type species: Acropelta omeiensis (C. Chr.) Nakai (= Polystichum TAXONOMICTREATMENT omeienseC. Chr.). PPoollyyssttiicchhuummloRnocthhi,tiTsen(tL..f)l.RoGtehrm(.=P3:ol3y1p,o6d9iu(m17l9o9n)c.hiTtyispeL.s).pecies: eParlneadcnttbstrotaesnrucrbheesetrdre;icatlr,aarnoerdlyempoisslttiotllhyiocnu,infbrearrraoenulscy;helddoiw,c-tolyreovcsertleeleeippci;inpghs,yetoterswd.ietRchhuimrzboooetmnset, Hypopeltis Michx., Fl. bor.-amer. 2: 266 (1803). Type species: closely towidely spacedpersistentstipebases, andpaleae. Fronds HypopeltislobulataBory (=Polystichumaculeatum(L.) Roth). monomorphic,caespitoseorcloselytowidelyspaced,to 1.8mlong: PlecosorusFee,Mem.foug.5: 150(1852).Typespecies:Plecosorus stipe proximally convex adaxially, becoming slightly to deeply mexicanusFee,nom.superfl.forCheilanthesspeciosissimaKunze sulcate distally; with two larger near-circular vascular bundles (= Polystichum speciosissimum (Kunze) R.M. Tryon & A.F. dorso-laterally,ventrallywiththreetofivesmallercircularvascular Tryon). bundles; initially moderately to densely paleated, becoming near Sorolepidium H. Christ inBot. Gaz. 51: 350 (1911). Type species: glabrouslater,thepaleaeoftenappearingheteromorphous,variable: Sorolepidium glaciale (H. Christ) H. Christ (= Polystichum lamina 1-pinnateto3-pinnate,anadromous,sometimesbearing 1 to glaciate H. Christ). severalpaleatedproliferousbudsadaxiallyalongtherachisnearthe ROUX 36 J.P. laminaapex:rachisadaxiallyshallowlytodeeplysulcate,thesulcus Distal pinnae neverfolded ventrally along the rachis; spores aborted proximally not open to sulci of lower order axes, moderately to 9.P.xsaltum denselypaleated;paleaevariable:pinnaeshort-stalked,oppositeto 10 Rhizometo 10mmindiameter;soriexindusiate 11 aplitnenrantaet,e,acclroossecloypitcoalwliydealuyricslpeadc:edp,inonfat-ernacimhbirsicaadtaex,iaslilmyplseultcoat2e,- Rhizomemorethan 10mmindiameter;soriindusiate 12 opentosulciofcostae,sparselytodenselypaleated;paleaevariable: u Rhizome to 5 mm in diameter; stipe and rachis paleae with long pinnules proximally mostly short-stalked, opposite to alternate, flagelliformoutgrowthsalongthemar^gin 10.^P.marionense cplionsneullyetmoowsitdleyllyonsgpearcetdh,anofttheenniemxbtriicnat2e-,pmthneatperooxrimmaolreacdriosssceocptiecd _ Rthhiinz-owamleletdoc,eQllmsmalo-nngditahmeemtearr;gisntjpe fachispa,1e1a.eP.traQnrskweiitehnosuet species, herbaceous to firmly coriaceous, inaequilateral, ovate to ovate-rhomboid or trullate, often somewhat falcate, mostly 12 Laminawithaproliferousbudalongtherachisneartheapex acroscopically auricled in 2-pinnate or more dissected species, 12.P.magm icum lobate,dentateorserrate,sharp-tippedoraristate;variouslypaleated. Laminawithoutproliferousbudsalongtherachis 13 aVtenthaetimoanrfgriene,wphiennnatsteelryilber,ainmcmheedr,seadnaodrrroamioseuds,.,Stoenr.mcii.nractuilanrg,neesasreno-r 13 oRuhtigzroomwethpsalaelaoengcotnhsepimcauroguisnl;ysrmuaglolseer,sotfitpeen,wriatchhiasfaenwdlpoinngnaf.-_i.rl.a,i.cfhoirsm tiallyunisenate,bornemediallyonunabbreviatedveinbranches,or paleae basally with short and/or ,ong flliform outgrowths along the near or at a vein ending of mostly anadromous vein branches: margin 13.P.zambesiacum sporangium with 8-(13)-30 indurated annulus cells; stalk with aglbasnednutloarrpcreelslesnto,rpeeglltaatned,ulmaors,tl3-yJsehpreirastiestebnetl,otwhecampasruglie:n vianrdiuosuisulJmy pcRauhlrievzaeoedmbeamsaparalgllieynawaeilntohotouucttognrssohpowrtithcsua;onud./ssolm.rya.llrlouengrgosusetn,iipmseoe,nsatrtalecyhowiusittgharnsodhwotrfp,htisnsntaar,l-aoirngaghctht.,iohser sculptured, with or without gland-like cells. Spores monolete, the margin 14 lmaoesstulryas2/o3mtoew3/h4aotftshpeinsuploorsee,leonfgttehn,ptehrefoprearties.poCrheriormreogsuloamrelynfuomldbeedr, 14 Proximal acroscopicpinnuleto22mmlong; largerstipebasepaleae n=41, 82, 164; 2n=82, 164, 328; apogamous 123, 246. oftenbicolorous r Proximalacroscopicpinnuleusuallymorethan22mmlong;largerstipe Key tO the African Species OfPolystictlUm (includ- basepaleaeneverbicolorous 16 ing Marion and Prince Edward Islands) 15 Rhizomeshort-decumbentwithcrowdedstipebases,closelybranched; 1 Lamina 1-pinnate(rarely 1-pinnate-pmnatifid) l.P.macleae apogamous(32sporespersporangium).... 14.P.monticola - Lamina2--pi,nnateto3-pmnate 2 - Rhizomedecumbent,stoloniferous;sexual(641s5p.oPr.esdrpearcsopmoornantgainuumm) 2 Rhizomeshort,erecttosuberect,mostlyunbranched 3 16 Pinnules maequilaterally ovate to narrowly trullate, to 60 x 13 mm; Rhizomeshort-decumbenttowidelycreeping,mostlybranched .... 10 sporangium stalk glandularor eglandular; indusium with or without unicellularthin-walledcellsalongthemargin 16.P.incongruum 3 Largerrhizomeandstipebasepaleaewithlongunisenatehairsalong themarginandsuperficially 2.P.luctuosum Pinnules inaequilaterally ovate, ovate-oblong, ovate-rhomboid or trullate,to50x19mm;sporangiumstalkandindusiumalwayseglandular Largerrhizomeandstipebasepaleaewithoutlongunisenatehairsalong ,7 p Dunoens themarginandsuperficially 4 4 Lamina with 1-3 paleated proliferous buds along the rachis nearthe 1. Polystichum macleae (Baker) Diels in Engl. & Prantl, Nat. laminaapex 5 Pflanzenfam. 1(4): 190(1902),asmacleanii.Type:SouthAfrica, - Laminawithoutproliferousbudsalongtherachis.... ... 6 GinolcdonvFiaellldisb'u,sMhcuLmeidais34-sDurbakBeonlsubser3g0e3n0p(rKolp-elec'tPoitlgyrpiem,'sdeRseisgt- 5 Pinnulemarginsobtuselyserratetocrenate,neveraristate natedbySchelpe&Anthony(1986);BOL!,SAM!-isolectotypes). 3.P.volkensii p- ^ Pinnuleauricleandapexaristate 4.P.kilimanjaricum Aspi.d..ium macl.eae Baker in Hook.\f.t,iIcon, pi.: t. i16/5i4 /(i1o8o8/6:)\, as 6 Apices ofpaleaeterminating in a short subulatecell ora small thin- macleaii. walledcell,themarginssetwithshortstraightand/orangularoutgrowths 7 Plantsterrestrial,epilithic,orrarelyepiphytic.Rhizomedecumbent, mm mm to 200 long x 20 in diameter, densely set with roots, Apicesofpaleaealwaysterminatinginanacicularcell,themarginsset persistentstipebases,andpaleae;paleaeferrugineous,membranous withlongstraightand/orlongtwistedemarginatetoforkedoutgrowths tQchartaceouSi narrowly ovate orlanceolamte, to7 x 2 mm. Fronds caespitose, to7perplant,arcuate,to 1.47 long: stipeproximally 7 Stipeandrachismoderatelypaleated;paleaemostlyflatorirregularly castaneous,stramineousdistally,adaxiallysulcate,to670mmlong folded 5.P.aculeatum x g mm jn diameter, proximally densely paleated; larger paleae Stipeandrachisdenselypaleated;paleaemostlyhelicallytwisted broadlyattached,concolorousorbicolorous,theconcolorouspaleae 6.P.setiferum ferrugineous, the bicolorous paleae centrally darkbrown orblack, 8 Conspicuouslylargerpaleaemostlyconfinedtothestipe,rugose ... ovate' narrowly ovate' ornarrowly oblong' cordate' entireorwith 7 P.transvaalense short and/or long marginal outgrowths proximally, the apex flagelliform, terminating in a long filiform cell oran oblong thin- :onspicuouslylargerstipepaleaeextendingtotherachis,neverrugose walledcell to37x8mm;smallerpaleaeconcolorous,ferrugineous, membranous, narrowlyovate, lanceolate, narrowlytriangular, nar- 9 Distalpinnaefoldedventrallyalongtherachis(conduplicate); spores rowlyoblongtoacicular,cordate,cordate-imbricate,orshort-stalked, notaborted 8.P.wilsonii proximallyeroseand/orwithlongtwisted,filiformoutgrowths,the POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 37 apex flagelliform, terminating in afiliform cell or an oblong thin- Havelock(CC),Schutte4(BOL);HavelockMine,Dyer51(NU).2632(Bela walled cell: lamina 1-pinnate (rarely 1-pinnate-pinnatifid), oblong Vista):Mbabane,NgwenyaMountain(AA),Compton31405(NBG) tonarrowlyelliptic,withupto37freepinnapairs,to840mmlong, WITHOUTEXACTLOCALITY:locoincerto,Boluss.n.(PRE);South the proximal pinnae slightly reduced, usually deflexed: rachis Africa, Woods.n.(NU). stramineous, adaxially sulcate, moderately paleated; paleae short- Thechangeofthespecificepithetmacleaiitomacleaeisinconcord- stalked,ferrugineous,membranoustochartaceous,narrowlyovate, ance with Article 60.11 (Recommendation 60C.l.a) of the narrowly lanceolate, oroblongto subulate,cordate, cordate-imbri- InternationalCodeofBotanicalNomenclature(Greuteretal.,1994). cate, proximally usually erose and/or with a few short or long twistedmarginaloutgrowths,theapexlong-attenuatetoflagelliform, DIAGNOSTICFEATURESANDRELATIONSHIPS. Polystichummacleae terminatinginanacicularcelloranoblongthin-walledcell,to3mm istheonly 1-pinnate(rarely 1-pinnate-pinnatifid) speciesinAfrica long: pinnae firmly herbaceous, olive-green adaxially, paler andisquitesimilartoP. kalambatitrenseTardieufromMadagascar. abaxially, generally not overlapping, short-stalked, narrowly ob- Baker (1886) considered P. macleae to be related to P. munitum blornoga-da-tctuenneuaattee,tsotrtariugnhcta,taeu,rdicouulbaltyeascerrroastceo,pitcoal1l6y8,mthmeblaosneguxne1q6uamllmy P(rKeasullff.r)oCm.MPardeesilraf.roTmhiNsoratshsuAmmpetiroincawaasndprPo.bfaabllcyinbealsleudmo(nSwt.h)eC1.- wide,theacroscopicauricleonproximalpinnaeoftenfree,ovateto pinnate lamina morphology in these taxa. Both these species, trullate, to 24 x 22 mm; costaadaxially sulcate, sparselypaleated, however,haveciliatedindusia.Also,thepaleamorphologyofboth the paleae taeniform, sessile or short-stalked, entire, the apex ter- these species differs from that in P. macleae. In P. macleae the minating in anacicularcelloranoblongthin-walledcell, to 3 mm marginal outgrowths ofthe lamina paleae are pluricellular whilst long,abaxiallymoderatelytodenselypaleated,thepaleaeferrugin- thoseofP. munitumandP.falcinellumareunicellular. eous, membranous, narrowly lanceolate to narrowlytrullate, often VARIATION. Polystichummacleaeshowsconsiderablevariationin bullate, cordate to cordate-imbricate, the margin proximally with thenumberofinduratedannuluscellspersporangiumandinindu- short and/or long irregular outgrowths, entire distally, the apex siumandpinnamorphology.Thenumberofinduratedannuluscells mtemrminatingin anacicularcelloranoblongthin-wmalmledcell,to2.4 persporangiumrangesfrom 12to28.Themeannumberofindurated long. Venation raised. Sori circular, to 1.5 in diameter, annuluscellspersporangiumis 16.67(n=650,SD=0.14)takenfrom variableinsize, thoseclosesttothecostalargest,discreteatmatu- 13 populations throughout the species' distribution. Some popul- rity, medial to inframedial on unabbreviated vein branches: ationshavealargernumberofinduratedannuluscellsthanothers. sporangium with 12-(16)-28 indurated annulus cells; stalk Althoughnodefinitecorrelationcouldbemadebetweenhabitatand eglandular:indusiumbrown,persistent,peltate,circulartoirregular, the number of indurated annulus cells, plants collected from an repand to erose, often with flabellate central processes, the maxi- exposedstreambankontheHartbeesvlaktenearLydenburg [Kluge mumradius0.29-(0.49)-0.7mm.Spores64persporangium,brown, 2039 (PRE)] have anumberofinduratedannuluscells thatranges theperisporefoldedtoformasparsereticulumoflowcompressed from 19 to 28 (x=23.86, SD=1.91, n=50), which is significantly ridges,variouslygranulate,verruculatetoechinulate,closelyperfo- higher than for a plant growing in a forest habitat [Thorncroft 68 rated, the exospore 40-(51.31)-66 x 28-(37.89)-48 urn. (NBG)], where the number of cells ranges between 12 and 16 Chromosomenumber2n=164. (x=13.28, SD=0.75, n=50). Intermediates between these extremes do occur. Indusia are mostly simple, buton some plants they may bear one or more small wings, whilst on others they may bear MATERIALEXAMINED SOUTHAFRICA. 2330 (Tzaneen): Tzaneen,WoodbushForestReserve numerousflabellatecentralprocesses.Themarginsvaryfromrepand M(uCCt)t,erB2al6s4i(nPhRasE)2;1W6o6od(bPRuEs)h;,WVoalnkJbaearrgs,vAelgda6t\ha10Fo(rBeOsLt).Re2s4e3rv0e(,Pi1l5g0r0imm',s toAeros1e-.pinnate-pinnatifid form of Polystichum macleae has been Rest): Haffenden Heights, Zoutpansberg (AA), Junod 4069 (P, PRE); recordedfromMpumalangawiththecentralpinnaebearingupto 11 Mariepskop (DB), Van der Schijf 4305, (B, NU, PRE), 5597 (PRE); nearly free pinnule pairs. Pinnules are inaequilaterally narrowly Mariepskopsummit,1800m,VanderSchijf'4861(PRE);Mariepskop,below trullate to oblong-attenuate in outline with the margins obtusely radarstation,Krynauw786(PRE); Graskop, Erasmus Kop,Hardcastle59 serrate.Inthedistalpinnaethemarginsaremerelylobedmidwayto (PRE);Mariepskop,Schweickerdt4305 (BOL);OhrigstadNatureReserve, thecosta.Soriareuniseriateoneithersideofthecostaandareborne 6000ft(DC),Jacobsen 1556 (PRE); Pilgrim's Rest, MountShebaNature inframedially. Reserve,Roux2555 (NBG);MountShebaNatureReserve,Jacobsen4436 Thesizeoftheacroscopicauriclevariesconsiderably,andonthe (SRcPohRclEki)e;(bDeMDno),u9n7R2ta5uShh(eP&RbESac)h;NlaiGterubaresenkRo9ep7s,4e4rDv(rePi,ReEkC)or;poGuGrcoahrsgk6eo3,p3,WK(aNogUwe)yr;n'1Gs7r3aPsa(skPsoR,pE,R)a;CuihPgial&r- paurroixcilmealsopmientniameeistiosvoefrtleanpsdewtiatchhetdhefrpoimnntahedriersetctolfythaebopvien.na.PiTnhnea grim'sRest,Rogers14925, 14927(PRE);BlydeBosboustasie,Bredenkamp marginsaregenerallyobtuselyserratedordoublyserratedbutrarely s.n.(PRE);Pilgrim'sRest,MacLea170(PRE);Kowyn'sPass,Schelpe1641 themarginsarealsodeeply lobedandserrated. (BOL,NH,NU),6092(BOL);Graskop,Fairyland,Roux2548,2549(NBG); DISTRIBUTIONANDECOLOGY. Polystichummacleaeisconfinedto GLHSarymradutsetbknseobe3pus8,rvglT(,ahPkRetHEeaP),ri;tn1nbS9aea6ceb0lsieevm,,l,a4fkM5otro0eeh0slt(fetBa,tA2B)T8r,wa8ei(KtePhflRwouEang)ite;teei2Pn2i0l03(g97Br,Bi()mB2,'O3sLW3R)a3e.gste2(,r5PMR35oE03)u(;(nLPtyLRdyAEedn)ned;bneSubrarusbgroi)gne:,,/ tophcrecouvDrirrniacnkegenoafstbSeeolreugvtahEtsiAcofnarsripcrmaaennagtinndagntdbheeWtownolerektnbheer1rgn350pinarattnshdeof1M9pS6uw0maazmil.laanTnghdae, Lydenburgroad,Roux2242, 2561 (NBG); Witklip Staatsbos (BD),Kluge speciesislargelyconfinedtoforestswhereitgrowsonbanksabove 806 (PRE); Belfast (CA), Wager s.n. (PRE); Kaapse Hoop (DB), Van streams, in forest margins, among rocks and often as a low-level Jaarsveld2Q8%a(NBG,PRE),3376(BOL);KaapseHoop,Wager73(BOL), epiphyte. Plants often form large stands in deep shade, but rarely 1496c(PRE). 2531 (Komatipoort): Barberton,TigerCreek,4500ft(CC), alsooccurinexposedhabitats. Thorncroft96(BR,P,Herb.PIC.SERM.,PRE);Barberton,MaidoftheMist, HGTahaovmrenelcorRcoekfs,te5r5v00e0,(0P,oftPn,RSfEca)hr,eml6Jp8oes(e4Nf1Bs1dG5a,l(,PB,1OP6LR4,E0)Pm;R,E1)K7;umniWil.tezso&fSHEOaotvtfeolBao1rc5bke,(Jr,StoPonRngEti)o.mwvaerldos 2. TPyolpyesatsicfohruAmslpuicdtiuuomsluumct(uKousnuzme)KuT.nzMeo.ore,Ind.fil.: 95 (1858). SWAZILAND. 2531 (Komatipoort): New Havelock, 12 miles from Fig. 3. ROUX 38 J.P. 10 mm Fig.2 Polystichummacleae.A,proximalpartoflamina;B,rhizome;C,sectionofabaxialsurfaceoffertilepinna.A&B,drawnfromRoux2548(NBG); C,drawnfromRoux2242(NBG). POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 39 Aspidium luctuosum Kunze in Linnaea 10: 548 (1836). Type: In angium with 10-(13)-19 induratedannuluscells; stalkeglandular: monte Katriviersberg in sylvis, Ecklon s.n. (LZf-syntype); ad indusium peltate, circular, entire, repand or crenulate, persistent, fontesfl. KatrivierpropePhilipstown,insylvismontium,Ecklon brown, pale brown and often dark centred before drying, cupulate s.n. (LZt-syntype). whendry,themaximumradius0.5-(0.73)-0.95 mm.Spores32per Aspidium tsus-simense Hook., Sp.fil. 4: 16, t. 220 (1862). Type: sporangium, brown, theperispore unevenly foldedto form narrow Island of Tsus Sima, in the Straits of Korea, Wilford s.n. (K- and broad reticulate ridges, the ridges and areas between ridges holotype, 2 sheets; NBG!-photograph). echinulate, spiculateorverruculate,theexospore30-(38.84)-50 x Polystichumtsus-simense (Hook.)J. Sm.,Hist,fil.: 219 (1875). 22-(28.2)-36 urn. Chromosomenumber2n=123, apogamous. Polystichum lobatum var. luctuosum (Kunze) H. Christ in Ber. MATERIALEXAMINED Schweiz. Bot. Ges. 3: 34(1893). LESOTHO. 2828 (Bethlehem): Leribe (CC), Phillips s.n. (SAM). 2927 (Maseru): RomaValley(BC), Schmitz6963 (PRE); Roma,Ruch 1909A- Plants terrestrial, epilithic, omrrmarely low-level epiphytes. Rhizome only(PRE). short,erecttosuberect,to10 indiameter,denselysetwithroots, SOUTHAFRICA.2430(Pilgrim'sRest):TheDowns(AA),Junods.n. persistent stipe bases, and paleae; larger paleae broadly attached, & 4044 (BR, P, PRE); Mt Sheba (DC), Kluge 2320 (NBG, PRE); Blyde castaneous, chartaceous, ovate, narrowlyovate, orlanceolate, cor- ForestReserve(DD),Jacobsen4365,4376(PRE);DriekopGorge,Graskop, date, with long twisted uniseriate, gland-tipped hairs on the apical Wager 178(PRE); Pilgrim'sRest,Ponies Krantz,Braithwaite229(BOL); marginandsurface,theapexflagelliform,terminatinginanoblong Sabie,just outside Ceylon Forest Reserve, Braithwaite 135 (BOL). 2530 thin-walled cell, to 10.5 x 3.3 mm; smaller paleae short-stalked, (Lydenburg): Lydenburg District, Spitzkop, Wilms 1781 (B, BM); Sabie narrowly triangular to subulate, cordate, the margins proximally G(oBOrLg)e;(SBaBb)i,e,WRaoggeerrss.2n.03(7P9RE()P;REL)o;neSuCdrweaelkaFCaalvless,,Sa1b5i0e0,mBu(rBrCo)w,sKl1u3g4e2 with numerous long and twisted uniseriate hairs, distally with 2463(NBG,PRE);Lydenburg,BuffelskloofNatureReserve,Burrows3860 widely spaced apically and basally directed marginal outgrowths (GRA);betweenMachadodorpandBadplaas(CD),Steel242(PRE).2531 thatbecomesmallerapically,theapexflagelliform,terminatingina (Komatipoort):RimersCreek,Barberton(CC),Thorncroft35(P);Baberton, smallthin-walledcell. Frondscrowded,caespitose,7-16perplant, Pott-Leendertz 5574 (PRE); Lomati Falls behind Barberton, Wager 154 m suberect to arching, to 0.93 long: stipe proximally castaneous, (PRE); Barberton, Thorncroft 36, 104c (PRE); creeks near Barberton, stramineous distally, adaxially sulcate, to450 mm long x 5 mm in Thorncroft2475(L).2729(Volksrust):Newcastle,NkanduReserve,4900ft cdaisatmaenteeoru,s,decnhsaertlaycepoaulse,atoevda;te,prcoorxdiamtael,ppraolxeiamealblryoeandtliyreaotrtawcihtehd,a ((DNDB)G,);SmPiotnhg6o4la(NBUu)s.h27N3at0u(rVeryRbeuserrgv)e:,ro1a5d00tomLu(nBeCb)u,rgGl(eAnD)2,3R9o0ux(P2R2E6)8; few short and/or long uniseriate hairs, distally with numerous JUtorhencshtto,nDeo2n9k6er(hNoUe)k.,25852080(fBt,eDthelveehneims)h:1F1a4r4m(PBRoEs)c;hkHllooobafn(eD,B)M,toRloauFxor1e2s2t8, multicellularhairsasforrhizomepaleae;distalpaleaeshort-stalked, (NBG); Witsieshoek,Junods.n. (P); Royal Natal National Park, Okell60 narrowlyoblong,narrowlytriangularorsubulate,cordatetohastate, (NU).2829(Harrismith): VanReenen,5000ft(AD),Schlechter6718(B, the margins bearing a few long and/or short multicellular hairs BM,GRA,PRE,SAM);VanReenensPass,Rehmann7204(P);VanReenen, proximally,distallywithwidelyandirregularly spacedoutgrowths 5000ft,Lidey42(NU);Robinson'sBush,OliviershoekPass(CA),Schelpe reduced in size and number towards apex, the apex flagelliform, 7967 (BOL); near Cathedral Peak, Box 3371 (BM). 2929 (Underberg): terminating in a small thin-walled cell, to 15 x 1.5 mm: lamina 2- Giants Castle Nature Reserve (AB), Roux 2503 (NBG); Injasuti Nature pinnateto2-pinnate-pinnatifid,withupto25freepinnapairs,to480 Reserve, below Cataract Valley, Roux 2718 (NBG); Champagne Castle, mm long, firmly herbaceous to coriaceous, olive-green adaxially, Bayer 1444 (NU); Cathedral Peak, bank of Kweliquala River, 4700 ft, prsuaelldceuarcteaedb,a,dxeioanflstleeyln,ynspaoarmlreeoawwthelday;topvdaaeltfeelaeteoxesohdvo:artter-,asctthahileskpersd,otrxdaiammriaknlebporiuonsw,nnaaetdosalxbiilgaahlctlkl,yy dmS(rcN,ahUlGe)olP;peeeatCkagP,th4heRkb(aiNeinUun)rPb;4aor5wCk5a,2GtohH(eroBdgwRrel,a,eltP5tRP5eE05a)03k;,f(Ct,5aN7tHCh0)oe0;wdrafEta,nsltKc9iPol6eulari(ktcN,,kUc)N1.;1o55l0C5ea0tm(0haNfeUtd,F,roMaraPleRcsPEtGe,)ra;e4k,2gC0ao1t0r5h45ef3t0-, glossy,chartaceoustocrustaceous,narrowlytriangulartosubulate, Edwards2685(NU,PRE);CathkinPeak,NdemaForest,4400ft,Hillary106 cordate to hastate, the auricles usually bearing long and twisted & 107 (NU); Cathkin Park, Howlett 53 (NH); Injasuti area, 5000 ft, multicellularanduniseriatehairssomeofwhichterminateinathin- Esterhuysen26034(BOL,NBG,PRE);aboveDaltonBridge,aboveBushmans walled cell, the margins either distally with short, widely and River,c.4500ft,Wrightetal.27(NH,PRE);MooiRiver,TheHoek,4700ft irregularly spaced outgrowths that reduce in size and number to- (BC), Bourquin 320 (NU); Polela District, Ndumduma, Glengariff(CB), wards the apex, or more or less entire in smaller paleae, to 7 mm Rennie913,940(NU);CobhamForestStation,WhaleRock,Hill48(GRA); ulopngt:op1i2nfnraeeepsihnonrtu-lsetaplaikresd,,n1a-rprionwnlaytelatnoce1o-lpaitnen,atper-opxininmaatlimlfyimdw,idweiltyh ABCllulalsrwokepsrpon8M3o19u1n(1tN(aUN)iH;n,n(eNDaUBr))B;,ulBVwuaelnrw,eJrSa,cahrSesulvnpseeeltdP,5652426(08N0U)(ft.N,BR2eG9n,3n0iP(eRPE5i)e4;t6e(rBNmuaUlr)wi;etrBzub(luDwrDge))r:,, spaced, distally often somewhat overlapping, to 173 long: Balgowan,farmBoschfontein,4000ft(AC),Fisher638(NH,NU),642(NH, pinna-rachisstramineous,adaxiallysulcate,denselysetwithpaleae NU);LionsRiverDistrict,Dargle,Smook624(NU);Balgowan,Thomas71 similartobutlesscomplexthanthoseontherachis:pinnuleswidely (NU); Balgowan,Devlin62 (NU); Balgowan,4000ft, Lindahl 107 (NU); spacedtooverlapping,theproximalacroscopicpinnulethelargest, BalgowanDistrict,Thienel109(NU);Balgowan,3500ft,Bernele113(NU); oftensignificantlylongerthanthenext,upto40mmlongand12mm Balgowan, 3500 ft, Crookes 105 (NU); Nottingham Road, McClean 899 wide, inaequilateral, narrowly trullate torhomboid, basiscopically (NH,PRE);NottinghamRoad,sinecoll.NH-26790(NH);Dargle,Griffin's cuneate,acroscopicallytruncateandauricled,oftensomewhatfalcate, Farm, 1500 m, Jones 20 (NH); Lions River, Dargle, Esterhuysen 26202 laocbraotsec-ospeircraaltley, ianrciisstaetde;toproorxniemaarllypitonntuhleescosshtoa;rt-csotsatlakeadd,axoifatlelny (RTiyBvsOeoLrn),;LsB.inao.lngs(oPBwRauEns),h;BFoPoisreecsthte,rHmoMaeorkil,tlz1b84u2r09g0,(m,Bc.OMLo2,l2l0N90U0)5f;t,(PBiCOeaLtre,nremNgaUir,ietPz6Rb9Eu2r)g;(L(NiCUoB)n);,s sulcate,glabrous,abaxiallysparselypaleated,thepaleaecastaneous, Pietermaritzburg,Blackridge,F.G.C.692(NU);Inanda,Woods.n.(B).3029 chartaceous, narrowly triangular-hastate to subulate-hastate, cor- (Kokstad): LangewachtForestReservenearKokstad,c. 1200m(CB),De datetocordate-imbricate,proximallywithlongand/orshortfiliform Joncheeres.n.(PRE);MtCurrie,Kokstad,Stephany505(BOL);GlenHope, outgrowthsoftenterminatinginathin-walledcell,theapexalways Jaconet & Jaconet 539 (BM). 3126 (Queenstown): Woodvale Forest, terminating in a small thin-walled cell, to 0.3 mm long. Venation Gwatyn,4200ft(AA), Galpin 8203 (PRE). 3127(LadyFrere): Engcobo immersed. Soricircular, c. 1.2 mm in diameter, terminal ornearly (DB); McLoughlin 1022 (PRE); Engcobo, Flanagan 2781 (PRE). 3128 terminalonabbreviatedveinbranches,essentiallyuniseriate:spor- (Umtata): Maclear, farm Woodcliffs (AB), Roux2479 (NBG); Engcobo, 40 J.P.ROUX 10mm Fig.3 Polystichumluctuosum.A,proximalpartoflamina;B,rhizome;C,abaxialsurfaceoffertilepinnule.AlldrawnfromRoux2433(NBG). POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 41 BESF(KaooAuArssD-Metc)Hs))h,tl:;b,oeCipHrinahangwets,skecyylahBlv6ziovi6ilniss0ugFssaoa(drd3B9e9O5pspLet(e()dBdB;(eOOeAmLTLCms)))om;,.mloooMnC3,nto2atiBu2iwsens6eltBeB(o1BoFFs1asooc1zcrrhihteb(ybseaBBte,reOr(gaLgBBu,)aC(f;)uoDM,rrATat)s6Cc)o,4a:lO4owBw,Kae(aarBGtk7)nxb.l3aee131rly28gi(28sBb.F45Oono.Lmr(()evBS(s;,uoGtmRNBFe(AqOorBr,aLsCed)e)sP;u,tt, sApbIlinlosadsmenuortesisita,vathaewers,iinhtavhutoaimwArobinfaa.ertirlIicanooarnfgnAefitprdhnliaedactugnartrnasaeptephpelaeodavafnairtvnnssangrtutihlloaeautrsibngoeeucnrmeleibilnnnesvdbriuporsertoirahannsmAgtpefehonrsatrinfaaclrAanlnsogyimiaaunni1mdn0pdsAltuhaoscnoiet1wads9s.n., AHodlalmasnd14s.2n.(N(UN)B;G)H;ogKsabtabcekrg,,MHaudtotnonnas.an.nd(BC,hLi)l;dKFaatllbser(gDBF)o;reGsrtse,atc.he3a0d0s0.nft., SwphoerreesastoionsAhsoiwanvaprliaanttison,thweitnhumAbseiranrpalnagnetss bheavtiwnegenlar1g0eransdpor2e1.s (SRGH);HogsbackForest,AucklandKloof,Griffenx46(PRE);Brambledene, than African plants (Table 1). Although variations in paleacolour Menziesberg,Acocks 11112 (PRE); Hogsback, Gibbs-Russel 3832 (PRE). occur,theirmorphologyremainsfairlystablethroughoutthedistri- 3227(Stutterheim): IsidingeForest(CA),Roux 1986(NBG);Keiskamma butionofthespecies. (HCoBe)k,,GRxouulxu2M4o3u3nt(aiNnB,G)5;50P0infte,SFtoorreyst3,50a9lo(nPgREA)m;atKoallaoghtraailF,orReostuxSta2t7i0o9n DISTRIBUTION AND ECOLOGY. In South Africa Polystichum (NBG);PirieForest,Flanagan 1758(PRE); FortCunningham,Roux2427 luctuosumoccursfromtheEasternCapethroughKwaZulu-Natalto (NBG); Pirie (CC), sine coll. s.n. (GRA); Amabele (DA), sine coll. s.n. the northeastern parts of the Free State, Mpumalanga, and the (PRE);Pirie,Sims.n.505,1727c(GRA,PRE);Komgha(DB),Flanagans.n. Northern Province. It also extends to the lower elevations in the (SASMW)A.ZILAND.2631(Mbabane):Gobolo,c. 3500ft(AC),Dlaminis.n. wwietshteirsnolpaatretdspoofpuLleastoitohnos,octcheurhriignhgera-lloynigntghepaeratstoefrnSweaszcialrapnmde,ntanidn G(wNeBZnGIi,,MNBMHtA,.BCPWaRsEEh):e;l,ISntCyrhaoanmsgaea,,1c.N0y84a30n0(g0aSnRftiG,,HC)6a.0m0p0ioftn,2C5h8a2s2e(3N8B1G3,(PNRUE,).SRGH); cZeinmtbraalbwpaer.tsOuotfsMiaddeaogfaAsfcrairc,aetxhteesnpdeicnigestooctchuersInodniaRneusnubicoonntainndentth,e WITHOUTEXACTLOCALITY:GoldFields,Ayress.n.(NH);Capde Pakistan,China,Vietnam,SouthKoreaandJapan(Honsyu,Sikoku BonneEsperance,Dreges.n. (P);Johannesburg, Westemans.n. (P);Capb. and Kyusyu). Spei,Ecklons.n.(P);Natal,sinecoll. s.n.(P);MorBridge,Hill692(PRE); Polystichumluctuosumoccursintheeasternpartsofthesummer Drakensberg,Bottomleys.n.(PRE);Natal,Buchanans.n.(BOL,M);O.F.S. rainfallregionsofsouthernAfricawhereitappearstoberestricted TM1761c(PRE);Natal,Wood504(PRE);invollbusmontiumseciisKatrivier, tothedrierforesttypessuchasDohneSourveldintheEasternCape, propePhilipstown,3000-4000ft,Ecklon&Zeyhers.n.(P);Natal,Buchanan Highland Sourveld along the Drakensberg foothills, 'Ngongoni 21A-only (M); Kaffrarianforests,Sim s.n. (B); Natalia, Buchanan74, 84 Veld in the KwaZulu-Natal midlands and Northeastern Mountain sH(K(iBaBin)t)mre;;aiclNCvoaaialetylpra.aT,l,Ps,3.rRn0.W.a0ovbN0o.iH-ds-4Vpa90s.l.,07ne08.Eyc,5(fktN,C(lUhDNoa)rHnn;e)j&ig;Xieu,bZmsee7.elyn0r.oh0iwe(0BrMF)fow;t3r,n8PeM.dsr6cato,Dlm(oiBR.,)ny;bcn.5rec0oSlefp0dlet0ei35,dft41,Dt8Are(rneB(rdgMNiteeU)or);rss;y.,Son.lonQouBucste.oh&nl.ilAnCe(fcnrBeoirMndct)leao;y,s, NrSSaooynuuagtrnipvgnaeagnlnsdifbreiirotngmo.Sc6wcI7uan0rzsiSmloaatunttedohl,e1vAa7afn4trdi0iocnmaas,lPoa.wnshglheuirtcghetheausaolssioun1wm8vZ2eoi5lcmdcmbu.aerssbcwaaetrpeomlneevnMatotiuaonnntds Barckley95(GRA);sinecoll.s.n.(L);locoincerto,sinecoll.s.n.NH-26468 Polystichumluctuosummostlygrowsonrocksalongstreams,but (NH). oftenalsoasalow-levelepiphyteinmoistforests. Plantsoftenalso growonrocksawayfromwaterand infairlydryconditions. The African Polystichum luctuosum (Kunze) T. Moore and the AsianP. tsus-simense(Hook.)J.Sm.havebeenconsideredeitheras 3. Polystichumvolkensii(Hieron.)C.Chr.,Indexfilic.:97(1905). TypeasforAspidium volkensiiHieron. distincttaxa(Mitui, 1965, 1968;Hirabayashi, 1969;Daigobo, 1973; Fig. 4. Nakaike, 1975;Gibby, 1985;Punethaetal., 1988)orassynonymous (Hope, 1902;HookerinHooker&Baker, 1868).Plantsoccurringin AspidiumvolkensiiHieron.inH.G.A.Engler,Pflanzenw.Ost-Afrikas: thewesternIndo-Himalayanmountainshavebeenascribedtoeither 86(1895).Type:Tanzania,anderoberenGrenzedesWaldesiiber P. tsus-simense (Dixit, 1983) or to P. luctuosum (Khullar, 1987; Kiboscho, 3000m, Volkens 1520(B!-holotype). Punethaetal., 1988).Fraser-Jenkins(inGibby, 1985)considersthe PolystichumbarbatumC.Chr.inNotizbl.Bot. Gart.Berlin-Dahlem two taxa as vicariants. I have studied material throughout the 9: 178 (1924). Type: Kenya, Mt. Aberdare, pr. Kinangop, regio distributionrangeofthesetaxaandfindthemtobeconspecific. Hagenia abyssinica, c. 3300 m, Rob. E. & Th. C.E. Fries 2735 DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of (K!-holotype; B!-isotype). mm Polystichum luctuosum is the olive-green colour of the adaxial Plants terrestrial. Rhizome short, erect, to 10 in diameter, surfaceofthelaminaandthedarkerveinsseeninlivingplants.Itis densely set with roots, persistent stipe bases, and paleae; paleae alsoseparatedfromothertaxaintheregionbytheusuallyverydark sessileorshort-stalked,ferrugineous,chartaceous,narrowlylanceo- rahnidznoamreraonwdpastliepaeeboacsceurprailnegaealboenagrtlhoengstfiipleifaonrdmroaucthgisr.owTthheslaalrognegr mlatme,lcoonrgd.atFer,oenndtsirec,aetshpeitaopseex, ttoer1m4inpaetrinpglainnt,anereaccti,cutloar1.c2elml, ltoon1g:5 athnedmtaarkgeionnaancduppaulleaatesufrofracme.whInednusmiaatuarree.laProgley,spteirscihsutmenltuacntduoesntuimrei,s tsoti5p2e0prmomximlaolnlgyxca1s0tmanmeoiunsd,isatmreatemri,nedoeunssedlisytaplallye,aatdeadx;ilaalrlgyersuplaclaetaee, furthermore atriploid apomict with 32 spores persporangium and short-stalked, ferrugineous, membranous, translucent, shrivelled, hasasomaticchromosomenumberof2n=123. elliptic to ovate, cordate to cordate-imbricate, minutely fimbriate, WithinthestudyareaPolystichumluctuosumistheonlymember theapexlong,shrivelled,filiform,entire,terminatinginanacicular belongingto sectionXiphopolystichum Daigobo. cell, to34x 10mm; smallerpaleaeshort-stalked,convolute,ovate to narrowly ovate, cordate to cordate-imbricate, proximally with VARIATION. Polystichum luctuosum showslittlevariation in stipe, short straight or angular marginal outgrowths, becoming entire laminaandbasalpinnalengthwithinthe studyarea. Acomparison towardstheapex,theapexsubulate,terminatinginanacicularcell: mm of these parts with Asian material shows that African (including lamina 3-pinnate, herbaceous, narrowly elliptic, to 925 long, Madagascar) plants are slightly larger than the plants from Asia. olive-greenadaxially,slightlypalerabaxially,withasinglepaleated Guard-cell length in African material shows little variation, but in proliferous bud on the rachis near the lamina apex, the proximal Asian plants the variation is pronounced. Asian plants also have pinnae decrescent, often somewhat deflexed: rachis stramineous, larger guard cells than African plants. Sori may be uniseriate or adaxiallysulcate,denselypaleated;paleaeshort-stalked,convolute, 42 J.P.ROUX Table1 VariationinmetriccharactersforAfricanandAsianPolystichumluctuosum(Kunze)T.Moore. Character

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The genus Polystichum (Dryopteridaceae) in Africa PDF

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Preview The genus Polystichum (Dryopteridaceae) in Africa

Bull. nat.Hist. Mus.Land. (Bot.)30(2): 33-79 Issued30November2000 The genus Polystichum (Dryopteridaceae) in " Africa ./ . \") , JACOBUS P. ROUX 22 NOV NationalBotanicalInstitute, ComptonHerbarium, PrivateBagX7, Claremont7735, SouthAfrica igfgKAL LJ6RARV CONTENTS Introduction 33 Materialsandmethods 34 Terminology 34 Taxonomictreatment 35 KeytotheAfricanspeciesofPolystichum(includingMarionandPrinceEdwardIslands) 36 References 78 Systematicindex 79 SYNOPSIS. PolystichumRothisaferngenusof160to200speciesoccurringthroughoutthetemperatepartsoftheworldandthe montanetropics,butabsentfromthearidregionsaswellasthelowlandtropics.AlthoughfloristicaccountsofPolystichumexist formanypartsoftheworld,thegenusremainspoorlyunderstoodandataxonomicaccountforthegroupasawholehasnever beenundertaken.PolystichumispoorlyrepresentedinAfrica(includingtheMarionIslandgroup)withmerely16speciesandone knownhybridhavingbeenrecordedfromtheregion.Mostofthespeciesareconfinedtosub-SaharanAfrica,occurringmainly alongtheeasternmountainrangesandMt. Camerooninthewest. InthisreviewoftheAfricanPolystichumspeciesdetailed observationsarepresentedespeciallyonthepaleae, asthese structureswerefoundtoprovidethebestcharactersonwhicha subgenericclassificationcanbebased.Diagnosticfeaturesandrelationships,variation,andthedistributionandecologyofeach speciesareincluded. INTRODUCTION allopolyploids (Vida& Reichstein, 1975; Wagner, 1979). Alsothe frequentoccurrenceofFl-hybridsinsomegroupsobscuresspecies limits. The genus Polystichum Roth consists of between 160 (Tryon & Aformalsubgenericclassificationforthegenushasthusfaronly Tryon, 1982)and200(Daigobo, 1972)species.Itoccursthroughout been provided for the east Asian species by Tagawa (1940) and thetemperatepartsoftheworldaswellasthemontanetropics,but Daigobo(1972).Someofthesesectionshavesincebeensubdivided is mostly absent from the lowland tropics. Within the range two further by Zhang & Kung (1995, 1996a, b) and Kung & Zhang distinctcentresofdiversitycanbeidentified,namelyalargerAsiatic (1998) to make provision for some ofthe Chinese species. Since centrewithapproximately70speciesandatropicalAmericancentre mostspeciesremainpoorlyknownaphylogenyforthegenuscannot with approximately 55 species. Both these regions fall within the beproposed. mountainoustropicscharacterizedbymild,moistclimatesthatare Althoughfloristicaccountsofthegenushavebeenpublishedfor hardly seasonal. They also correspond with the areas of highest manypartsoftheworld,somebeingveryoldandoutdated,nosingle speciesdiversityforhomosporousferns (Tryon, 1985). monographictreatmentexistsandmostspeciesremainpoorlyknown. WithintheDryopterideaePolystichumismostcloselyrelatedto Some ofthe modern-day regional treatments provide no detailed Arachniodes Blume, Cyrtomium C. Presl, Dryopteris Adans. and observations that may suggest affinities. Within the study area Phanerophlebia C. Presl. Cyrtomium and Phanerophlebia have floristic accounts for Polystichum are available for North Africa often been included in Polystichum (Kramer, 1990), but it has (Maire, 1952), West tropical Africa (Alston, 1959), Cameroon subsequently been shown that both these genera have a closer (Tardieu-Blot, 1964), Mozambique, Malawi, Zambia and Zimba- affinitytoPolystichumthantoeachother(Yatskievych, 1996). bwe (Schelpe, 1970), Rwanda, Burundi and Kivu (Democratic Polystichuminastrictsenseisanaturalandrelativelyhomogene- RepublicofCongo)(PichiSermolli,1985),southernAfrica(Schelpe ousgroupofplantscharacterizedby 1-pinnateto3-pinnatelaminae &Anthony, 1986) andBioko(Benl, 1991). withacroscopicallydevelopedultimatesegments,anadromousfree It is a well-known fact that Africa, when compared with other venation, uni- orbiseriatecircularsori positioned medially, termi- tropical parts of the world, supports a floristically impoverished nally or near terminally on abbreviated or unabbreviated vein vascularflora.Thisphenomenonisalsoreflectedinthepteridophyte branches,andpeltateindusia(anumberofspeciesareexindusiate). floraofthecontinent.Thecauseofthisfloristicpovertyisascribed Thereceptacleappearstobenudeinmostspecies. to the isolation ofAfricafrom the othercontinents since the mid- Although Polystichum as a genus is easily recognized, many Cretaceousandthesubsequentsignificantchangesintheclimateas speciesaresuperficiallyverysimilar.Thedelimitationofspeciesis aresultofuplift,continentaldriftandaridificationcausedbyextra- furtherhamperedbythefactthatmany,mostlycommonspeciesare tropical glaciation. All these changes may well have resulted in a TheNaturalHistoryMuseum,2000 34 J.P.ROUX progressive elimination of the once rich tropical and subtropical mountain systems, whilstP. transvaalense and P. wilsonii are dis- foreststhatexistedtowardsthelateJurassicandtheestablishmentof tributedthroughoutseven mountainsystems.ThedistributionofP. extensivedeserts and semi-desertsbytheearly Pliocene (Coetzee, wilsonii, however, also extends along the Himalaya mountains to 1993). Bhutan, Japan and Taiwan. Polystichum luctuosum has an almost Within the study area two regions can be identified: an African similareastern distribution butis confinedtotheDrakensberg and and a sub-Antarctic region. The origin, composition and floristic Chimanimani mountain systems in Africa. Polystichum luctuosum affinitiesoftheseregionsdiffermarkedly.TheAfricanregionisthe andP.wilsoniialsoshowadisjunctdistributionwiththeMadagascan largest and today there are three fundamentally different floras or region. biogeographical subregions which can be identified: a southern Basedonobservationstaken fromthesystematic treatment, and Africanflora,atropical AfricanfloraandaNorthAfricanflora. judging from wide-ranging species, southern and tropical African The flora ofthe southern African subregion is believed to have PolystichumhasacloseraffinitywithtaxafromAsiathanwiththose evolved gradually since the mid-Tertiary, derived partly from an from the Americas. Polystichum in the North African phyto- ancientsouthernAfricantemperatefloraandpartlyfromatropical geographical subregion shows a closer affinity with Polystichum African forest flora (Goldblatt, 1978). Elements of the southern fromEuropethanfromAfricaasP. aculeatumandP. setiferumare AfricanfloracurrentlyextendintotropicalAfricaalongtheeastern widespreadinthatregion.OnlyP. marionense,endemictoMarion, escarpment.ThesouthernCapeforestsarebelievedtobeimpover- Prince Edward and Crozet Islands, occurs in the sub-Antarctic & ished remnants of the tropical African forest flora (Coetzee phytogeographical region. Muller, 1984). Also the tropical African flora is believed to be an impoverishedremnantofaoncemuchrichertropicalrainforestflora thatextendedoverafargreaterareathanitcurrentlyoccupies. MATERIALSANDMETHODS North Africa has also experienced significant changes in its climate and vegetation. During the Palaeocene the present Sahara desert was clothed by a rich tropical lowland rainforest that also This review is basedon observations made duringextensive field- coveredpartofEurope(Greenway, 1973;Raven&Axelrod, 1974), work in southern Africa and on cultivated plants collected during butbytheOligo-Mioceneitwasreplacedbyasubtropicalwoodland thesetravels.Thecollectionsofseveralherbariawerealsostudied. cslaivmaantneaes(tAaxbellirsohded&itsRelafveinn,the19m7a8)j.orFpraortmotfhethePlSiaohcaernae (aQudeezseelr,t TNhHe,seNUin,clPu,dPe:REB,,BRMA,B,BOSLA,M,BRS,REGTHH,,WGRAAG,(Ka,bLb,reMv,iaMtiAoLns,fNoBllGo,w 1978), serving as an effective barrierto migration from the south. Holmgren et al., 1990) and the private herbarium ofProf. R.E.G. The formation of glaciers on the high mountains during the Pichi Sermolli (PIC.SERM.). Pleistocene permittedtheestablishmentofcircumborealelements. Paleaandindusiumobservationsweremadebyremovingasmall ManyoftheseelementsarepresentintheNorthAfricanflorasince number of these structures from selected specimens. These were it is composed of relict elements of African origin as well as cleaned and cleared in diluted household bleach, after which they elements from Eurasia, not frequent in the present sub-Saharan were semi-permanently mounted in glycerine and the cover slips flora. The mediterranean influence on the flora of North Africa sealed with Entellan. Observations were made with an Olympus justifies it being considered as a biogeographical subregion ofits CH-2light-microscopefittedwithadrawingtube. own. Thecollections studiedareall listedunder 'Materialexamined'. The Marion and Prince Edward Island group forms part ofthe These are arranged alphabetically according to country oforigin. sub-Antarcticregion,aphytogeographicalareacompletelydifferent South African (including Lesotho and Swaziland) collections are from the foregoing. This island group is ofvolcanic origin and is furtherarr&angedaccordingtothequarter-degreesquare-gridsystem estimatedtobe0.5 millionyearsold(Verwoerd, 1971). Situatedin (Edwards Leistner, 1971).Inthissystemeachone-degreesquare theSouthernOceansome 1800kmfromAfrica,itsbiotaconsistsof is known by a standardized name, derived from a town or other ttahxeAambscoeaulptvaebtslwoei-nothfhailbroidtnsagot-sfditnshotetaAnafclrewiadcyiassnpfepratsveaorluirdaaonbpdlheytthfeeosraabprilelainlttiymgitrtooewedstithna.btlhiesihr dfaeinavdtiuDdreedfroionftmoilmfepoftuorrttohaarnlicfgeh-tdeiagnnrdetehteospqsutqaouraberoset.(t3oE0m'a.cxEh3a0co')hn,eh-nadulemfg-brdeeeregerdeseqAus,aqrBue,arCies ohcocwuervreern,cearteoctlhoesecloyntrielnaetnetd(tKoortanxaas,in1e9i9t3h)e.rTthroepimcaajloArimteyriocfathaensde/, nisumagbaeirnedsuAb,diBv,iCdeadndinDt.oBqyuaurstienrg-dtehgerseeecos-qouradriensat(e1s5'axgeo1g5'r)a,phaigcaailn orsoutheastAsia.Pteridophytesofthesub-Saharabiogeographical areacanimmediatelybeidentified. regionexhibitthreediscontinuousdistributionpatterns: an Ameri- Unlesscitedotherwise, thechromosomenumbersprovidedhere can-Africandisjunction,anAfrican-Madagascandisjunctionandan are based on the author's own observations and will be published African-Asiandisjunction. elsewhere. Polystichum in Africa is largely confined to the Afromontane Phytochorion. White (1978)dividedthismontanearchipelagointo Terminology sevenregional mountain systems. TheNorth African Atlas moun- This study is principallybasedon adetailedcomparative morpho- tain ranges arehereaddedasaneighth. Althoughthe sub-Saharan logicalanalysisofthe sporophyte, wherepaleastructureprovedto mountainrangesaresufficientlydistinct,thesystemsareconnected bemostinformativeinsuggestingspeciesgroups.Thetermsusedto byacomplexseriesofintermediatefloras(Fig. 1).TheDrakensberg describetheapexortheapicalcellofthepaleaearedefinedas: system, with six Polystichumendemics, is the richest. This is also true for the angiosperms (White, 1978). The only other mountain - apexflagelliform:theapexofthepaleaterminatesinauniseriate systemswithtrueendemicsaretheImatongs-Usambarasystemwith seriesofslendercells. twoendemics(P. kilimanjaricum&P. volkensii)andtheEthiopian - subulatecell:theapicalcellislessthan0.4mmlongandtheapex systemwithoneendemic(P.magnificum).OtherAfricanPolystichum isusuallyblunt. specieshavewiderdistributions.Polystichumzambesiacumoccurs - acicularcell:theapicalcellisslender,straight,morethan0.4mm in the Chimanimani, Uluguru-Mulanje, and Imatongs-Usambara long, andtheapex isusually sharp. POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 35 V r VIII. NORTHAFRICAN 1 IV. IMATONGS-USAMBARA U-y\P.kilimanjaricum VII. ORAKENSBERG '.drakensbergense .incongruum macleae . .monticola .pungens .transkeiense Fig. 1 DistributionofPolystichuminAfrica. filiformcell:theapicalcellisslender,twisted,andmorethan0.6 HemesteumH.Lev.,Fl.Kouy-Tcheou:450,496(1915),nonNewm. mm long. (1851). Type species: severalPolystichumspeciesarelisted. thin-walledcell:thesecellscanvaryconsiderablyinsizeandthe Aetopteron HouseinAmer. FernJ. 10: 88 (1920), nom. nud. cell wall is conspicuously thinner than that ofthe surrounding Papuapteris C. Chr. in Brittonia 2: 300 (1937). Type species: cells, and is therefore often lost. In dry material the cellular Papuapteris linearis C. Chr. (= Polystichum lineare (C. Chr.) contents areusuallyyellowishincolourandappearcrystalline. Copel.). Acropelta Nakai in Bull. Natl. Sci. Mus. Tokyo33: 5 (1953). Type species: Acropelta omeiensis (C. Chr.) Nakai (= Polystichum TAXONOMICTREATMENT omeienseC. Chr.). PPoollyyssttiicchhuummloRnocthhi,tiTsen(tL..f)l.RoGtehrm(.=P3:ol3y1p,o6d9iu(m17l9o9n)c.hiTtyispeL.s).pecies: eParlneadcnttbstrotaesnrucrbheesetrdre;icatlr,aarnoerdlyempoisslttiotllhyiocnu,infbrearrraoenulscy;helddoiw,c-tolyreovcsertleeleeippci;inpghs,yetoterswd.ietRchhuimrzboooetmnset, Hypopeltis Michx., Fl. bor.-amer. 2: 266 (1803). Type species: closely towidely spacedpersistentstipebases, andpaleae. Fronds HypopeltislobulataBory (=Polystichumaculeatum(L.) Roth). monomorphic,caespitoseorcloselytowidelyspaced,to 1.8mlong: PlecosorusFee,Mem.foug.5: 150(1852).Typespecies:Plecosorus stipe proximally convex adaxially, becoming slightly to deeply mexicanusFee,nom.superfl.forCheilanthesspeciosissimaKunze sulcate distally; with two larger near-circular vascular bundles (= Polystichum speciosissimum (Kunze) R.M. Tryon & A.F. dorso-laterally,ventrallywiththreetofivesmallercircularvascular Tryon). bundles; initially moderately to densely paleated, becoming near Sorolepidium H. Christ inBot. Gaz. 51: 350 (1911). Type species: glabrouslater,thepaleaeoftenappearingheteromorphous,variable: Sorolepidium glaciale (H. Christ) H. Christ (= Polystichum lamina 1-pinnateto3-pinnate,anadromous,sometimesbearing 1 to glaciate H. Christ). severalpaleatedproliferousbudsadaxiallyalongtherachisnearthe ROUX 36 J.P. laminaapex:rachisadaxiallyshallowlytodeeplysulcate,thesulcus Distal pinnae neverfolded ventrally along the rachis; spores aborted proximally not open to sulci of lower order axes, moderately to 9.P.xsaltum denselypaleated;paleaevariable:pinnaeshort-stalked,oppositeto 10 Rhizometo 10mmindiameter;soriexindusiate 11 aplitnenrantaet,e,acclroossecloypitcoalwliydealuyricslpeadc:edp,inonfat-ernacimhbirsicaadtaex,iaslilmyplseultcoat2e,- Rhizomemorethan 10mmindiameter;soriindusiate 12 opentosulciofcostae,sparselytodenselypaleated;paleaevariable: u Rhizome to 5 mm in diameter; stipe and rachis paleae with long pinnules proximally mostly short-stalked, opposite to alternate, flagelliformoutgrowthsalongthemar^gin 10.^P.marionense cplionsneullyetmoowsitdleyllyonsgpearcetdh,anofttheenniemxbtriicnat2e-,pmthneatperooxrimmaolreacdriosssceocptiecd _ Rthhiinz-owamleletdoc,eQllmsmalo-nngditahmeemtearr;gisntjpe fachispa,1e1a.eP.traQnrskweiitehnosuet species, herbaceous to firmly coriaceous, inaequilateral, ovate to ovate-rhomboid or trullate, often somewhat falcate, mostly 12 Laminawithaproliferousbudalongtherachisneartheapex acroscopically auricled in 2-pinnate or more dissected species, 12.P.magm icum lobate,dentateorserrate,sharp-tippedoraristate;variouslypaleated. Laminawithoutproliferousbudsalongtherachis 13 aVtenthaetimoanrfgriene,wphiennnatsteelryilber,ainmcmheedr,seadnaodrrroamioseuds,.,Stoenr.mcii.nractuilanrg,neesasreno-r 13 oRuhtigzroomwethpsalaelaoengcotnhsepimcauroguisnl;ysrmuaglolseer,sotfitpeen,wriatchhiasfaenwdlpoinngnaf.-_i.rl.a,i.cfhoirsm tiallyunisenate,bornemediallyonunabbreviatedveinbranches,or paleae basally with short and/or ,ong flliform outgrowths along the near or at a vein ending of mostly anadromous vein branches: margin 13.P.zambesiacum sporangium with 8-(13)-30 indurated annulus cells; stalk with aglbasnednutloarrpcreelslesnto,rpeeglltaatned,ulmaors,tl3-yJsehpreirastiestebnetl,otwhecampasruglie:n vianrdiuosuisulJmy pcRauhlrievzaeoedmbeamsaparalgllieynawaeilntohotouucttognrssohpowrtithcsua;onud./ssolm.rya.llrlouengrgosusetn,iipmseoe,nsatrtalecyhowiusittgharnsodhwotrfp,htisnsntaar,l-aoirngaghctht.,iohser sculptured, with or without gland-like cells. Spores monolete, the margin 14 lmaoesstulryas2/o3mtoew3/h4aotftshpeinsuploorsee,leonfgttehn,ptehrefoprearties.poCrheriormreogsuloamrelynfuomldbeedr, 14 Proximal acroscopicpinnuleto22mmlong; largerstipebasepaleae n=41, 82, 164; 2n=82, 164, 328; apogamous 123, 246. oftenbicolorous r Proximalacroscopicpinnuleusuallymorethan22mmlong;largerstipe Key tO the African Species OfPolystictlUm (includ- basepaleaeneverbicolorous 16 ing Marion and Prince Edward Islands) 15 Rhizomeshort-decumbentwithcrowdedstipebases,closelybranched; 1 Lamina 1-pinnate(rarely 1-pinnate-pmnatifid) l.P.macleae apogamous(32sporespersporangium).... 14.P.monticola - Lamina2--pi,nnateto3-pmnate 2 - Rhizomedecumbent,stoloniferous;sexual(641s5p.oPr.esdrpearcsopmoornantgainuumm) 2 Rhizomeshort,erecttosuberect,mostlyunbranched 3 16 Pinnules maequilaterally ovate to narrowly trullate, to 60 x 13 mm; Rhizomeshort-decumbenttowidelycreeping,mostlybranched .... 10 sporangium stalk glandularor eglandular; indusium with or without unicellularthin-walledcellsalongthemargin 16.P.incongruum 3 Largerrhizomeandstipebasepaleaewithlongunisenatehairsalong themarginandsuperficially 2.P.luctuosum Pinnules inaequilaterally ovate, ovate-oblong, ovate-rhomboid or trullate,to50x19mm;sporangiumstalkandindusiumalwayseglandular Largerrhizomeandstipebasepaleaewithoutlongunisenatehairsalong ,7 p Dunoens themarginandsuperficially 4 4 Lamina with 1-3 paleated proliferous buds along the rachis nearthe 1. Polystichum macleae (Baker) Diels in Engl. & Prantl, Nat. laminaapex 5 Pflanzenfam. 1(4): 190(1902),asmacleanii.Type:SouthAfrica, - Laminawithoutproliferousbudsalongtherachis.... ... 6 GinolcdonvFiaellldisb'u,sMhcuLmeidais34-sDurbakBeonlsubser3g0e3n0p(rKolp-elec'tPoitlgyrpiem,'sdeRseisgt- 5 Pinnulemarginsobtuselyserratetocrenate,neveraristate natedbySchelpe&Anthony(1986);BOL!,SAM!-isolectotypes). 3.P.volkensii p- ^ Pinnuleauricleandapexaristate 4.P.kilimanjaricum Aspi.d..ium macl.eae Baker in Hook.\f.t,iIcon, pi.: t. i16/5i4 /(i1o8o8/6:)\, as 6 Apices ofpaleaeterminating in a short subulatecell ora small thin- macleaii. walledcell,themarginssetwithshortstraightand/orangularoutgrowths 7 Plantsterrestrial,epilithic,orrarelyepiphytic.Rhizomedecumbent, mm mm to 200 long x 20 in diameter, densely set with roots, Apicesofpaleaealwaysterminatinginanacicularcell,themarginsset persistentstipebases,andpaleae;paleaeferrugineous,membranous withlongstraightand/orlongtwistedemarginatetoforkedoutgrowths tQchartaceouSi narrowly ovate orlanceolamte, to7 x 2 mm. Fronds caespitose, to7perplant,arcuate,to 1.47 long: stipeproximally 7 Stipeandrachismoderatelypaleated;paleaemostlyflatorirregularly castaneous,stramineousdistally,adaxiallysulcate,to670mmlong folded 5.P.aculeatum x g mm jn diameter, proximally densely paleated; larger paleae Stipeandrachisdenselypaleated;paleaemostlyhelicallytwisted broadlyattached,concolorousorbicolorous,theconcolorouspaleae 6.P.setiferum ferrugineous, the bicolorous paleae centrally darkbrown orblack, 8 Conspicuouslylargerpaleaemostlyconfinedtothestipe,rugose ... ovate' narrowly ovate' ornarrowly oblong' cordate' entireorwith 7 P.transvaalense short and/or long marginal outgrowths proximally, the apex flagelliform, terminating in a long filiform cell oran oblong thin- :onspicuouslylargerstipepaleaeextendingtotherachis,neverrugose walledcell to37x8mm;smallerpaleaeconcolorous,ferrugineous, membranous, narrowlyovate, lanceolate, narrowlytriangular, nar- 9 Distalpinnaefoldedventrallyalongtherachis(conduplicate); spores rowlyoblongtoacicular,cordate,cordate-imbricate,orshort-stalked, notaborted 8.P.wilsonii proximallyeroseand/orwithlongtwisted,filiformoutgrowths,the POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 37 apex flagelliform, terminating in afiliform cell or an oblong thin- Havelock(CC),Schutte4(BOL);HavelockMine,Dyer51(NU).2632(Bela walled cell: lamina 1-pinnate (rarely 1-pinnate-pinnatifid), oblong Vista):Mbabane,NgwenyaMountain(AA),Compton31405(NBG) tonarrowlyelliptic,withupto37freepinnapairs,to840mmlong, WITHOUTEXACTLOCALITY:locoincerto,Boluss.n.(PRE);South the proximal pinnae slightly reduced, usually deflexed: rachis Africa, Woods.n.(NU). stramineous, adaxially sulcate, moderately paleated; paleae short- Thechangeofthespecificepithetmacleaiitomacleaeisinconcord- stalked,ferrugineous,membranoustochartaceous,narrowlyovate, ance with Article 60.11 (Recommendation 60C.l.a) of the narrowly lanceolate, oroblongto subulate,cordate, cordate-imbri- InternationalCodeofBotanicalNomenclature(Greuteretal.,1994). cate, proximally usually erose and/or with a few short or long twistedmarginaloutgrowths,theapexlong-attenuatetoflagelliform, DIAGNOSTICFEATURESANDRELATIONSHIPS. Polystichummacleae terminatinginanacicularcelloranoblongthin-walledcell,to3mm istheonly 1-pinnate(rarely 1-pinnate-pinnatifid) speciesinAfrica long: pinnae firmly herbaceous, olive-green adaxially, paler andisquitesimilartoP. kalambatitrenseTardieufromMadagascar. abaxially, generally not overlapping, short-stalked, narrowly ob- Baker (1886) considered P. macleae to be related to P. munitum blornoga-da-tctuenneuaattee,tsotrtariugnhcta,taeu,rdicouulbaltyeascerrroastceo,pitcoal1l6y8,mthmeblaosneguxne1q6uamllmy P(rKeasullff.r)oCm.MPardeesilraf.roTmhiNsoratshsuAmmpetiroincawaasndprPo.bfaabllcyinbealsleudmo(nSwt.h)eC1.- wide,theacroscopicauricleonproximalpinnaeoftenfree,ovateto pinnate lamina morphology in these taxa. Both these species, trullate, to 24 x 22 mm; costaadaxially sulcate, sparselypaleated, however,haveciliatedindusia.Also,thepaleamorphologyofboth the paleae taeniform, sessile or short-stalked, entire, the apex ter- these species differs from that in P. macleae. In P. macleae the minating in anacicularcelloranoblongthin-walledcell, to 3 mm marginal outgrowths ofthe lamina paleae are pluricellular whilst long,abaxiallymoderatelytodenselypaleated,thepaleaeferrugin- thoseofP. munitumandP.falcinellumareunicellular. eous, membranous, narrowly lanceolate to narrowlytrullate, often VARIATION. Polystichummacleaeshowsconsiderablevariationin bullate, cordate to cordate-imbricate, the margin proximally with thenumberofinduratedannuluscellspersporangiumandinindu- short and/or long irregular outgrowths, entire distally, the apex siumandpinnamorphology.Thenumberofinduratedannuluscells mtemrminatingin anacicularcelloranoblongthin-wmalmledcell,to2.4 persporangiumrangesfrom 12to28.Themeannumberofindurated long. Venation raised. Sori circular, to 1.5 in diameter, annuluscellspersporangiumis 16.67(n=650,SD=0.14)takenfrom variableinsize, thoseclosesttothecostalargest,discreteatmatu- 13 populations throughout the species' distribution. Some popul- rity, medial to inframedial on unabbreviated vein branches: ationshavealargernumberofinduratedannuluscellsthanothers. sporangium with 12-(16)-28 indurated annulus cells; stalk Althoughnodefinitecorrelationcouldbemadebetweenhabitatand eglandular:indusiumbrown,persistent,peltate,circulartoirregular, the number of indurated annulus cells, plants collected from an repand to erose, often with flabellate central processes, the maxi- exposedstreambankontheHartbeesvlaktenearLydenburg [Kluge mumradius0.29-(0.49)-0.7mm.Spores64persporangium,brown, 2039 (PRE)] have anumberofinduratedannuluscells thatranges theperisporefoldedtoformasparsereticulumoflowcompressed from 19 to 28 (x=23.86, SD=1.91, n=50), which is significantly ridges,variouslygranulate,verruculatetoechinulate,closelyperfo- higher than for a plant growing in a forest habitat [Thorncroft 68 rated, the exospore 40-(51.31)-66 x 28-(37.89)-48 urn. (NBG)], where the number of cells ranges between 12 and 16 Chromosomenumber2n=164. (x=13.28, SD=0.75, n=50). Intermediates between these extremes do occur. Indusia are mostly simple, buton some plants they may bear one or more small wings, whilst on others they may bear MATERIALEXAMINED SOUTHAFRICA. 2330 (Tzaneen): Tzaneen,WoodbushForestReserve numerousflabellatecentralprocesses.Themarginsvaryfromrepand M(uCCt)t,erB2al6s4i(nPhRasE)2;1W6o6od(bPRuEs)h;,WVoalnkJbaearrgs,vAelgda6t\ha10Fo(rBeOsLt).Re2s4e3rv0e(,Pi1l5g0r0imm',s toAeros1e-.pinnate-pinnatifid form of Polystichum macleae has been Rest): Haffenden Heights, Zoutpansberg (AA), Junod 4069 (P, PRE); recordedfromMpumalangawiththecentralpinnaebearingupto 11 Mariepskop (DB), Van der Schijf 4305, (B, NU, PRE), 5597 (PRE); nearly free pinnule pairs. Pinnules are inaequilaterally narrowly Mariepskopsummit,1800m,VanderSchijf'4861(PRE);Mariepskop,below trullate to oblong-attenuate in outline with the margins obtusely radarstation,Krynauw786(PRE); Graskop, Erasmus Kop,Hardcastle59 serrate.Inthedistalpinnaethemarginsaremerelylobedmidwayto (PRE);Mariepskop,Schweickerdt4305 (BOL);OhrigstadNatureReserve, thecosta.Soriareuniseriateoneithersideofthecostaandareborne 6000ft(DC),Jacobsen 1556 (PRE); Pilgrim's Rest, MountShebaNature inframedially. Reserve,Roux2555 (NBG);MountShebaNatureReserve,Jacobsen4436 Thesizeoftheacroscopicauriclevariesconsiderably,andonthe (SRcPohRclEki)e;(bDeMDno),u9n7R2ta5uShh(eP&RbESac)h;NlaiGterubaresenkRo9ep7s,4e4rDv(rePi,ReEkC)or;poGuGrcoahrsgk6eo3,p3,WK(aNogUwe)yr;n'1Gs7r3aPsa(skPsoR,pE,R)a;CuihPgial&r- paurroixcilmealsopmientniameeistiosvoefrtleanpsdewtiatchhetdhefrpoimnntahedriersetctolfythaebopvien.na.PiTnhnea grim'sRest,Rogers14925, 14927(PRE);BlydeBosboustasie,Bredenkamp marginsaregenerallyobtuselyserratedordoublyserratedbutrarely s.n.(PRE);Pilgrim'sRest,MacLea170(PRE);Kowyn'sPass,Schelpe1641 themarginsarealsodeeply lobedandserrated. (BOL,NH,NU),6092(BOL);Graskop,Fairyland,Roux2548,2549(NBG); DISTRIBUTIONANDECOLOGY. Polystichummacleaeisconfinedto GLHSarymradutsetbknseobe3pus8,rvglT(,ahPkRetHEeaP),ri;tn1nbS9aea6ceb0lsieevm,,l,a4fkM5otro0eeh0slt(fetBa,tA2B)T8r,wa8ei(KtePhflRwouEang)ite;teei2Pn2i0l03(g97Br,Bi()mB2,'O3sLW3R)a3e.gste2(,r5PMR35oE03)u(;(nLPtyLRdyAEedn)ned;bneSubrarusbgroi)gne:,,/ tophcrecouvDrirrniacnkegenoafstbSeeolreugvtahEtsiAcofnarsripcrmaaennagtinndagntdbheeWtownolerektnbheer1rgn350pinarattnshdeof1M9pS6uw0maazmil.laanTnghdae, Lydenburgroad,Roux2242, 2561 (NBG); Witklip Staatsbos (BD),Kluge speciesislargelyconfinedtoforestswhereitgrowsonbanksabove 806 (PRE); Belfast (CA), Wager s.n. (PRE); Kaapse Hoop (DB), Van streams, in forest margins, among rocks and often as a low-level Jaarsveld2Q8%a(NBG,PRE),3376(BOL);KaapseHoop,Wager73(BOL), epiphyte. Plants often form large stands in deep shade, but rarely 1496c(PRE). 2531 (Komatipoort): Barberton,TigerCreek,4500ft(CC), alsooccurinexposedhabitats. Thorncroft96(BR,P,Herb.PIC.SERM.,PRE);Barberton,MaidoftheMist, HGTahaovmrenelcorRcoekfs,te5r5v00e0,(0P,oftPn,RSfEca)hr,eml6Jp8oes(e4Nf1Bs1dG5a,l(,PB,1OP6LR4,E0)Pm;R,E1)K7;umniWil.tezso&fSHEOaotvtfeolBao1rc5bke,(Jr,StoPonRngEti)o.mwvaerldos 2. TPyolpyesatsicfohruAmslpuicdtiuuomsluumct(uKousnuzme)KuT.nzMeo.ore,Ind.fil.: 95 (1858). SWAZILAND. 2531 (Komatipoort): New Havelock, 12 miles from Fig. 3. ROUX 38 J.P. 10 mm Fig.2 Polystichummacleae.A,proximalpartoflamina;B,rhizome;C,sectionofabaxialsurfaceoffertilepinna.A&B,drawnfromRoux2548(NBG); C,drawnfromRoux2242(NBG). POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 39 Aspidium luctuosum Kunze in Linnaea 10: 548 (1836). Type: In angium with 10-(13)-19 induratedannuluscells; stalkeglandular: monte Katriviersberg in sylvis, Ecklon s.n. (LZf-syntype); ad indusium peltate, circular, entire, repand or crenulate, persistent, fontesfl. KatrivierpropePhilipstown,insylvismontium,Ecklon brown, pale brown and often dark centred before drying, cupulate s.n. (LZt-syntype). whendry,themaximumradius0.5-(0.73)-0.95 mm.Spores32per Aspidium tsus-simense Hook., Sp.fil. 4: 16, t. 220 (1862). Type: sporangium, brown, theperispore unevenly foldedto form narrow Island of Tsus Sima, in the Straits of Korea, Wilford s.n. (K- and broad reticulate ridges, the ridges and areas between ridges holotype, 2 sheets; NBG!-photograph). echinulate, spiculateorverruculate,theexospore30-(38.84)-50 x Polystichumtsus-simense (Hook.)J. Sm.,Hist,fil.: 219 (1875). 22-(28.2)-36 urn. Chromosomenumber2n=123, apogamous. Polystichum lobatum var. luctuosum (Kunze) H. Christ in Ber. MATERIALEXAMINED Schweiz. Bot. Ges. 3: 34(1893). LESOTHO. 2828 (Bethlehem): Leribe (CC), Phillips s.n. (SAM). 2927 (Maseru): RomaValley(BC), Schmitz6963 (PRE); Roma,Ruch 1909A- Plants terrestrial, epilithic, omrrmarely low-level epiphytes. Rhizome only(PRE). short,erecttosuberect,to10 indiameter,denselysetwithroots, SOUTHAFRICA.2430(Pilgrim'sRest):TheDowns(AA),Junods.n. persistent stipe bases, and paleae; larger paleae broadly attached, & 4044 (BR, P, PRE); Mt Sheba (DC), Kluge 2320 (NBG, PRE); Blyde castaneous, chartaceous, ovate, narrowlyovate, orlanceolate, cor- ForestReserve(DD),Jacobsen4365,4376(PRE);DriekopGorge,Graskop, date, with long twisted uniseriate, gland-tipped hairs on the apical Wager 178(PRE); Pilgrim'sRest,Ponies Krantz,Braithwaite229(BOL); marginandsurface,theapexflagelliform,terminatinginanoblong Sabie,just outside Ceylon Forest Reserve, Braithwaite 135 (BOL). 2530 thin-walled cell, to 10.5 x 3.3 mm; smaller paleae short-stalked, (Lydenburg): Lydenburg District, Spitzkop, Wilms 1781 (B, BM); Sabie narrowly triangular to subulate, cordate, the margins proximally G(oBOrLg)e;(SBaBb)i,e,WRaoggeerrss.2n.03(7P9RE()P;REL)o;neSuCdrweaelkaFCaalvless,,Sa1b5i0e0,mBu(rBrCo)w,sKl1u3g4e2 with numerous long and twisted uniseriate hairs, distally with 2463(NBG,PRE);Lydenburg,BuffelskloofNatureReserve,Burrows3860 widely spaced apically and basally directed marginal outgrowths (GRA);betweenMachadodorpandBadplaas(CD),Steel242(PRE).2531 thatbecomesmallerapically,theapexflagelliform,terminatingina (Komatipoort):RimersCreek,Barberton(CC),Thorncroft35(P);Baberton, smallthin-walledcell. Frondscrowded,caespitose,7-16perplant, Pott-Leendertz 5574 (PRE); Lomati Falls behind Barberton, Wager 154 m suberect to arching, to 0.93 long: stipe proximally castaneous, (PRE); Barberton, Thorncroft 36, 104c (PRE); creeks near Barberton, stramineous distally, adaxially sulcate, to450 mm long x 5 mm in Thorncroft2475(L).2729(Volksrust):Newcastle,NkanduReserve,4900ft cdaisatmaenteeoru,s,decnhsaertlaycepoaulse,atoevda;te,prcoorxdiamtael,ppraolxeiamealblryoeandtliyreaotrtawcihtehd,a ((DNDB)G,);SmPiotnhg6o4la(NBUu)s.h27N3at0u(rVeryRbeuserrgv)e:,ro1a5d00tomLu(nBeCb)u,rgGl(eAnD)2,3R9o0ux(P2R2E6)8; few short and/or long uniseriate hairs, distally with numerous JUtorhencshtto,nDeo2n9k6er(hNoUe)k.,25852080(fBt,eDthelveehneims)h:1F1a4r4m(PBRoEs)c;hkHllooobafn(eD,B)M,toRloauFxor1e2s2t8, multicellularhairsasforrhizomepaleae;distalpaleaeshort-stalked, (NBG); Witsieshoek,Junods.n. (P); Royal Natal National Park, Okell60 narrowlyoblong,narrowlytriangularorsubulate,cordatetohastate, (NU).2829(Harrismith): VanReenen,5000ft(AD),Schlechter6718(B, the margins bearing a few long and/or short multicellular hairs BM,GRA,PRE,SAM);VanReenensPass,Rehmann7204(P);VanReenen, proximally,distallywithwidelyandirregularly spacedoutgrowths 5000ft,Lidey42(NU);Robinson'sBush,OliviershoekPass(CA),Schelpe reduced in size and number towards apex, the apex flagelliform, 7967 (BOL); near Cathedral Peak, Box 3371 (BM). 2929 (Underberg): terminating in a small thin-walled cell, to 15 x 1.5 mm: lamina 2- Giants Castle Nature Reserve (AB), Roux 2503 (NBG); Injasuti Nature pinnateto2-pinnate-pinnatifid,withupto25freepinnapairs,to480 Reserve, below Cataract Valley, Roux 2718 (NBG); Champagne Castle, mm long, firmly herbaceous to coriaceous, olive-green adaxially, Bayer 1444 (NU); Cathedral Peak, bank of Kweliquala River, 4700 ft, prsuaelldceuarcteaedb,a,dxeioanflstleeyln,ynspaoarmlreeoawwthelday;topvdaaeltfeelaeteoxesohdvo:artter-,asctthahileskpersd,otrxdaiammriaknlebporiuonsw,nnaaetdosalxbiilgaahlctlkl,yy dmS(rcN,ahUlGe)olP;peeeatCkagP,th4heRkb(aiNeinUun)rPb;4aor5wCk5a,2GtohH(eroBdgwRrel,a,eltP5tRP5eE05a)03k;,f(Ct,5aN7tHCh0)oe0;wdrafEta,nsltKc9iPol6eulari(ktcN,,kUc)N1.;1o55l0C5ea0tm(0haNfeUtd,F,roMaraPleRcsPEtGe,)ra;e4k,2gC0ao1t0r5h45ef3t0-, glossy,chartaceoustocrustaceous,narrowlytriangulartosubulate, Edwards2685(NU,PRE);CathkinPeak,NdemaForest,4400ft,Hillary106 cordate to hastate, the auricles usually bearing long and twisted & 107 (NU); Cathkin Park, Howlett 53 (NH); Injasuti area, 5000 ft, multicellularanduniseriatehairssomeofwhichterminateinathin- Esterhuysen26034(BOL,NBG,PRE);aboveDaltonBridge,aboveBushmans walled cell, the margins either distally with short, widely and River,c.4500ft,Wrightetal.27(NH,PRE);MooiRiver,TheHoek,4700ft irregularly spaced outgrowths that reduce in size and number to- (BC), Bourquin 320 (NU); Polela District, Ndumduma, Glengariff(CB), wards the apex, or more or less entire in smaller paleae, to 7 mm Rennie913,940(NU);CobhamForestStation,WhaleRock,Hill48(GRA); ulopngt:op1i2nfnraeeepsihnonrtu-lsetaplaikresd,,n1a-rprionwnlaytelatnoce1o-lpaitnen,atper-opxininmaatlimlfyimdw,idweiltyh ABCllulalsrwokepsrpon8M3o19u1n(1tN(aUN)iH;n,n(eNDaUBr))B;,ulBVwuaelnrw,eJrSa,cahrSesulvnpseeeltdP,5652426(08N0U)(ft.N,BR2eG9n,3n0iP(eRPE5i)e4;t6e(rBNmuaUlr)wi;etrBzub(luDwrDge))r:,, spaced, distally often somewhat overlapping, to 173 long: Balgowan,farmBoschfontein,4000ft(AC),Fisher638(NH,NU),642(NH, pinna-rachisstramineous,adaxiallysulcate,denselysetwithpaleae NU);LionsRiverDistrict,Dargle,Smook624(NU);Balgowan,Thomas71 similartobutlesscomplexthanthoseontherachis:pinnuleswidely (NU); Balgowan,Devlin62 (NU); Balgowan,4000ft, Lindahl 107 (NU); spacedtooverlapping,theproximalacroscopicpinnulethelargest, BalgowanDistrict,Thienel109(NU);Balgowan,3500ft,Bernele113(NU); oftensignificantlylongerthanthenext,upto40mmlongand12mm Balgowan, 3500 ft, Crookes 105 (NU); Nottingham Road, McClean 899 wide, inaequilateral, narrowly trullate torhomboid, basiscopically (NH,PRE);NottinghamRoad,sinecoll.NH-26790(NH);Dargle,Griffin's cuneate,acroscopicallytruncateandauricled,oftensomewhatfalcate, Farm, 1500 m, Jones 20 (NH); Lions River, Dargle, Esterhuysen 26202 laocbraotsec-ospeircraaltley, ianrciisstaetde;toproorxniemaarllypitonntuhleescosshtoa;rt-csotsatlakeadd,axoifatlelny (RTiyBvsOeoLrn),;LsB.inao.lngs(oPBwRauEns),h;BFoPoisreecsthte,rHmoMaeorkil,tlz1b84u2r09g0,(m,Bc.OMLo2,l2l0N90U0)5f;t,(PBiCOeaLtre,nremNgaUir,ietPz6Rb9Eu2r)g;(L(NiCUoB)n);,s sulcate,glabrous,abaxiallysparselypaleated,thepaleaecastaneous, Pietermaritzburg,Blackridge,F.G.C.692(NU);Inanda,Woods.n.(B).3029 chartaceous, narrowly triangular-hastate to subulate-hastate, cor- (Kokstad): LangewachtForestReservenearKokstad,c. 1200m(CB),De datetocordate-imbricate,proximallywithlongand/orshortfiliform Joncheeres.n.(PRE);MtCurrie,Kokstad,Stephany505(BOL);GlenHope, outgrowthsoftenterminatinginathin-walledcell,theapexalways Jaconet & Jaconet 539 (BM). 3126 (Queenstown): Woodvale Forest, terminating in a small thin-walled cell, to 0.3 mm long. Venation Gwatyn,4200ft(AA), Galpin 8203 (PRE). 3127(LadyFrere): Engcobo immersed. Soricircular, c. 1.2 mm in diameter, terminal ornearly (DB); McLoughlin 1022 (PRE); Engcobo, Flanagan 2781 (PRE). 3128 terminalonabbreviatedveinbranches,essentiallyuniseriate:spor- (Umtata): Maclear, farm Woodcliffs (AB), Roux2479 (NBG); Engcobo, 40 J.P.ROUX 10mm Fig.3 Polystichumluctuosum.A,proximalpartoflamina;B,rhizome;C,abaxialsurfaceoffertilepinnule.AlldrawnfromRoux2433(NBG). POLYSTICHUM(DRYOPTERIDACEAE)INAFRICA 41 BESF(KaooAuArssD-Metc)Hs))h,tl:;b,oeCipHrinahangwets,skecyylahBlv6ziovi6ilniss0ugFssaoa(drd3B9e9O5pspLet(e()dBdB;(eOOeAmLTLCms)))om;,.mloooMnC3,nto2atiBu2iwsens6eltBeB(o1BoFFs1asooc1zcrrhihteb(ybseaBBte,reOr(gaLgBBu,)aC(f;)uoDM,rrATat)s6Cc)o,4a:lO4owBw,Kae(aarBGtk7)nxb.l3aee131rly28gi(28sBb.F45Oono.Lmr(()evBS(s;,uoGtmRNBFe(AqOorBr,aLsCed)e)sP;u,tt, sApbIlinlosadsmenuortesisita,vathaewers,iinhtavhutoaimwArobinfaa.ertirlIicanooarnfgnAefitprdhnliaedactugnartrnasaeptephpelaeodavafnairtvnnssangrtutihlloaeautrsibngoeeucnrmeleibilnnnesvdbriuporsertoirahannsmAgtpefehonrsatrinfaaclrAanlnsogyimiaaunni1mdn0pdsAltuhaoscnoiet1wads9s.n., AHodlalmasnd14s.2n.(N(UN)B;G)H;ogKsabtabcekrg,,MHaudtotnonnas.an.nd(BC,hLi)l;dKFaatllbser(gDBF)o;reGsrtse,atc.he3a0d0s0.nft., SwphoerreesastoionsAhsoiwanvaprliaanttison,thweitnhumAbseiranrpalnagnetss bheavtiwnegenlar1g0eransdpor2e1.s (SRGH);HogsbackForest,AucklandKloof,Griffenx46(PRE);Brambledene, than African plants (Table 1). Although variations in paleacolour Menziesberg,Acocks 11112 (PRE); Hogsback, Gibbs-Russel 3832 (PRE). occur,theirmorphologyremainsfairlystablethroughoutthedistri- 3227(Stutterheim): IsidingeForest(CA),Roux 1986(NBG);Keiskamma butionofthespecies. (HCoBe)k,,GRxouulxu2M4o3u3nt(aiNnB,G)5;50P0infte,SFtoorreyst3,50a9lo(nPgREA)m;atKoallaoghtraailF,orReostuxSta2t7i0o9n DISTRIBUTION AND ECOLOGY. In South Africa Polystichum (NBG);PirieForest,Flanagan 1758(PRE); FortCunningham,Roux2427 luctuosumoccursfromtheEasternCapethroughKwaZulu-Natalto (NBG); Pirie (CC), sine coll. s.n. (GRA); Amabele (DA), sine coll. s.n. the northeastern parts of the Free State, Mpumalanga, and the (PRE);Pirie,Sims.n.505,1727c(GRA,PRE);Komgha(DB),Flanagans.n. Northern Province. It also extends to the lower elevations in the (SASMW)A.ZILAND.2631(Mbabane):Gobolo,c. 3500ft(AC),Dlaminis.n. wwietshteirsnolpaatretdspoofpuLleastoitohnos,octcheurhriignhgera-lloynigntghepaeratstoefrnSweaszcialrapnmde,ntanidn G(wNeBZnGIi,,MNBMHtA,.BCPWaRsEEh):e;l,ISntCyrhaoanmsgaea,,1c.N0y84a30n0(g0aSnRftiG,,HC)6a.0m0p0ioftn,2C5h8a2s2e(3N8B1G3,(PNRUE,).SRGH); cZeinmtbraalbwpaer.tsOuotfsMiaddeaogfaAsfcrairc,aetxhteesnpdeicnigestooctchuersInodniaRneusnubicoonntainndentth,e WITHOUTEXACTLOCALITY:GoldFields,Ayress.n.(NH);Capde Pakistan,China,Vietnam,SouthKoreaandJapan(Honsyu,Sikoku BonneEsperance,Dreges.n. (P);Johannesburg, Westemans.n. (P);Capb. and Kyusyu). Spei,Ecklons.n.(P);Natal,sinecoll. s.n.(P);MorBridge,Hill692(PRE); Polystichumluctuosumoccursintheeasternpartsofthesummer Drakensberg,Bottomleys.n.(PRE);Natal,Buchanans.n.(BOL,M);O.F.S. rainfallregionsofsouthernAfricawhereitappearstoberestricted TM1761c(PRE);Natal,Wood504(PRE);invollbusmontiumseciisKatrivier, tothedrierforesttypessuchasDohneSourveldintheEasternCape, propePhilipstown,3000-4000ft,Ecklon&Zeyhers.n.(P);Natal,Buchanan Highland Sourveld along the Drakensberg foothills, 'Ngongoni 21A-only (M); Kaffrarianforests,Sim s.n. (B); Natalia, Buchanan74, 84 Veld in the KwaZulu-Natal midlands and Northeastern Mountain sH(K(iBaBin)t)mre;;aiclNCvoaaialetylpra.aT,l,Ps,3.rRn0.W.a0ovbN0o.iH-ds-4Vpa90s.l.,07ne08.Eyc,5(fktN,C(lUhDNoa)rHnn;e)j&ig;Xieu,bZmsee7.elyn0r.oh0iwe(0BrMF)fow;t3r,n8PeM.dsr6cato,Dlm(oiBR.,)ny;bcn.5rec0oSlefp0dlet0ei35,dft41,Dt8Are(rneB(rdgMNiteeU)or);rss;y.,Son.lonQouBucste.oh&nl.ilAnCe(fcnrBeoirMndct)leao;y,s, NrSSaooynuuagtrnipvgnaeagnlnsdifbreiirotngmo.Sc6wcI7uan0rzsiSmloaatunttedohl,e1vAa7afn4trdi0iocnmaas,lPoa.wnshglheuirtcghetheausaolssioun1wm8vZ2eoi5lcmdcmbu.aerssbcwaaetrpeomlneevnMatotiuaonnntds Barckley95(GRA);sinecoll.s.n.(L);locoincerto,sinecoll.s.n.NH-26468 Polystichumluctuosummostlygrowsonrocksalongstreams,but (NH). oftenalsoasalow-levelepiphyteinmoistforests. Plantsoftenalso growonrocksawayfromwaterand infairlydryconditions. The African Polystichum luctuosum (Kunze) T. Moore and the AsianP. tsus-simense(Hook.)J.Sm.havebeenconsideredeitheras 3. Polystichumvolkensii(Hieron.)C.Chr.,Indexfilic.:97(1905). TypeasforAspidium volkensiiHieron. distincttaxa(Mitui, 1965, 1968;Hirabayashi, 1969;Daigobo, 1973; Fig. 4. Nakaike, 1975;Gibby, 1985;Punethaetal., 1988)orassynonymous (Hope, 1902;HookerinHooker&Baker, 1868).Plantsoccurringin AspidiumvolkensiiHieron.inH.G.A.Engler,Pflanzenw.Ost-Afrikas: thewesternIndo-Himalayanmountainshavebeenascribedtoeither 86(1895).Type:Tanzania,anderoberenGrenzedesWaldesiiber P. tsus-simense (Dixit, 1983) or to P. luctuosum (Khullar, 1987; Kiboscho, 3000m, Volkens 1520(B!-holotype). Punethaetal., 1988).Fraser-Jenkins(inGibby, 1985)considersthe PolystichumbarbatumC.Chr.inNotizbl.Bot. Gart.Berlin-Dahlem two taxa as vicariants. I have studied material throughout the 9: 178 (1924). Type: Kenya, Mt. Aberdare, pr. Kinangop, regio distributionrangeofthesetaxaandfindthemtobeconspecific. Hagenia abyssinica, c. 3300 m, Rob. E. & Th. C.E. Fries 2735 DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of (K!-holotype; B!-isotype). mm Polystichum luctuosum is the olive-green colour of the adaxial Plants terrestrial. Rhizome short, erect, to 10 in diameter, surfaceofthelaminaandthedarkerveinsseeninlivingplants.Itis densely set with roots, persistent stipe bases, and paleae; paleae alsoseparatedfromothertaxaintheregionbytheusuallyverydark sessileorshort-stalked,ferrugineous,chartaceous,narrowlylanceo- rahnidznoamreraonwdpastliepaeeboacsceurprailnegaealboenagrtlhoengstfiipleifaonrdmroaucthgisr.owTthheslaalrognegr mlatme,lcoonrgd.atFer,oenndtsirec,aetshpeitaopseex, ttoer1m4inpaetrinpglainnt,anereaccti,cutloar1.c2elml, ltoon1g:5 athnedmtaarkgeionnaancduppaulleaatesufrofracme.whInednusmiaatuarree.laProgley,spteirscihsutmenltuacntduoesntuimrei,s tsoti5p2e0prmomximlaolnlgyxca1s0tmanmeoiunsd,isatmreatemri,nedoeunssedlisytaplallye,aatdeadx;ilaalrlgyersuplaclaetaee, furthermore atriploid apomict with 32 spores persporangium and short-stalked, ferrugineous, membranous, translucent, shrivelled, hasasomaticchromosomenumberof2n=123. elliptic to ovate, cordate to cordate-imbricate, minutely fimbriate, WithinthestudyareaPolystichumluctuosumistheonlymember theapexlong,shrivelled,filiform,entire,terminatinginanacicular belongingto sectionXiphopolystichum Daigobo. cell, to34x 10mm; smallerpaleaeshort-stalked,convolute,ovate to narrowly ovate, cordate to cordate-imbricate, proximally with VARIATION. Polystichum luctuosum showslittlevariation in stipe, short straight or angular marginal outgrowths, becoming entire laminaandbasalpinnalengthwithinthe studyarea. Acomparison towardstheapex,theapexsubulate,terminatinginanacicularcell: mm of these parts with Asian material shows that African (including lamina 3-pinnate, herbaceous, narrowly elliptic, to 925 long, Madagascar) plants are slightly larger than the plants from Asia. olive-greenadaxially,slightlypalerabaxially,withasinglepaleated Guard-cell length in African material shows little variation, but in proliferous bud on the rachis near the lamina apex, the proximal Asian plants the variation is pronounced. Asian plants also have pinnae decrescent, often somewhat deflexed: rachis stramineous, larger guard cells than African plants. Sori may be uniseriate or adaxiallysulcate,denselypaleated;paleaeshort-stalked,convolute, 42 J.P.ROUX Table1 VariationinmetriccharactersforAfricanandAsianPolystichumluctuosum(Kunze)T.Moore. Character

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