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rc0OZ)Z6Zegzt(del Dt slr.nN Nr)tsid Vol. 13, No. 2 (2000)

Materials and methods

This study is basedon examination of herbarium collections from PERTH, as well as selected material from CANB, along with observationsmade on live of all speciesexcepl B. tubulosa. As only a singleAustraliancollection of B. tubulosowasavailable,the descriptionofthis taxonis panly basedon SouthAfrican materialhoused at NBG (seeHolmgren et al. 1990). All measurementswere madefrom herbariummaterial (reconstituted where necessary). Terminology for indumentumfollows Hewson( 1988).

Taxonomictr€atment

Synonymydata presented in thispaper is restrictedto namesthat have been applied to in Australia,or namesused by Lewis (1959)in her monographof the . For additional .nomenclaturalinformation and further synonymy refer to Phillips(1951) and Lewis (1959).

BabianaKer Gawl., Curtis's Bot. Mag., 15: sub t. 539( 1802).

Perennialherbs. Corzs usuallydeep-seated, tunics papery, decaying into fine or coarsefibres. S/emsubterranean or aerial,erect to declinateor decumbent,simple or sometimesbranched, terete to angular.Leavesbasal, distichous, usually plicate, sometimes tortuous, curled or undulate,lamina flat or rarelyterete, hairy or rarelyglabrous. a spike, flowers distichous, spiral or secund. Bractsand bracteoler sheathing, free or partiallyunited, herbaceous, apical portion usually scarious, hdry or glabrous.Flowers usually zygomorphic and bitabiate, sometimes actinomorphic and funnel- shaped,mostly shades of bluebut rarelyred, yellow or cream,often fragrant. Perianth tube straight or curved,cylindrical or funnel-shaped. Tepals equal or unequal.S/azeirs inserted in themouth of the pedanthtube, usually unilateral and arcuate, rarely syrnrnetdcally aranged and erect,anthers basifixed.Ovary glabrous or hairy.Sryle exserted or rarelyincluded; style branches 3, slenderor apicallyexpanded. Frrrit a cartilaginouscapsule, glabrous or hairy. Seedsseveral per locule, funicle prominentand swollen, rugose, shiny. Chromosome number x=7.

A genusof c. 65 speciesin southernAfrica with onein Socotra.

'baviaan' 'babianer', Etymology.DerivedfromtheDutch forbaboon oritsCape corruption thecommon nameapplied to Babianaspecies in the Caperegion of SouthAfrica, asa resultof their statusas a favouredfood of theseanimals (Lewis 1959).

Key to naturalizedlaxa of Babianaoccurring in WesternAustralia

1 Perianthtube more than twice as long asthe tepals,45-85mm long, flowerscream 1.B. tubulosavar. tubiflora l. Perianthtube shorter than to aslong asthe tepals, 7-18 mm long;flowers predominantlyblue to magenta 2. Bracteolesunited c. two-thirdsto four-fifthsof theirlength, herbaceous to themargins; flowers with themarked tepals lowermost; ovary glabrous or pubescenton theribs only ...... 2. B. nana 2. Bracteolesfree to thebase, mostly herbaceous, with a scarious,reddish-brown marginto 0.4 mm wide;flowers twisted such that the marked tepals are uppermostand facing the stemapex; ovary densely sericeous ...... 3. B' angustifolia '( 's:ed,{.raq8ray 'Supuedxe sercedsaerql eq1;6 ururoc 1'9) ,{1,tro1slreqlu srputr (6661 .{1n1lu se) tuetxo IIns sI ell€rlsnv ualsoi& ur uoqelndod u,trourl,{1uo aqJ eler1snv tuelse.d\ur pezrl€Jnleuserceds DuotqDg Jer.Ifoqlfti pesnJuoceq o1 ,41a1qunsr puE uox€l e^rlcuqsrp€ sr osoryqu DuorqDg 'sa:rN

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'Suruurw .fg V !{ ut ouDqDg Jo sercadspezlFmleu eqt Jo .^{ar^ercnuouoxEt V J I ? rqcsdel 286 NuytsiaYol. 13, No. 2 (2000) of Babiana nal\ralized in Westem Australia,B. tubulosais probably the least significant in terms of weediness,at leaston the basisofpresent observations(G.J. Keighery pers.comm.).

2. Babiananana (Andr.)Spreng., Syst. Veg. 1: 156(1825).

Illustrqtions.Jeppe, Spring and Winter FloweringBulbs of the Cape96-97 (1989)lManning & Goldblatt,West Coast: S. African Wild FlowerGuide 7: 70-71(1996); Hussey e/ al., Westem Weeds 29 (1997)las B. distichal.

Perennial herb 70-30 cm high. Co,'rnsubglobose to broadly ovoid, 10-20 mm in diameter,tunics papery,decaying into fine fibres. Stez 45-100 mm tong, cormlets occasionallypresent in axils of the subteffaneanportion, teaves distichous, spreading (fanwise) to erect, scarcely plicate, pubescent;sheathing base 20-80 mm long; lamina nafiowly obovate-eltipticto narrowly elliptic, 45-160 mm Iong, 7-25 mm wide, baseattenuate-oblique, apex acuminateto acute. Inflorescence shofier than to (usually) longer than ,1- or 2-branched,2-10-flowered, flowers not twisted to face the stem apex, straight to occasionallyflexuose, pubescent. Bracts sheathing,herbaceous, reddish-brown,t narrowly ovatein lateralview (marginsoften t straigho, 12-20 mm long, pubescent to sericeous-pubescent,scarious apical portion glabrous; apex acute to obtuse, rarely apiculate. Brqcteolesrr'itedfor c. two-thirds to four-fifths oftheir length,sheathing, herbaceous, reddish-brown, : narrowly ovate in lateral view (marginsoften t straight), l2-20 mm long, pubescentto sericeous- pubescent,scarious apical portion glabrousl apex of individual bracteolesacute to narowly acute. Perianth marv e to purpleor bluish-purple,the lower threetepals each with a whitish orcream areanear the middle andareddish mark nearthebase,apiculum ofouter tepalspurplish-black, glabrous;perianth tube staight, cylindrical proximally, flared towards the apex, 12-17 mm long; tepals subequal, narrowly obovate to obovate,25-40 mm long,6-10 mm wide; baseattenuate; apex obtuseto acute or the outer tepals apiculate. Stamensarching, filaments 10-14 mm long, anthersnarowly oblong, creamtomauve,5-5.5 mm long. Ovaryellipsoidto rarelyovoid,3-5 mm long,glabrous orpubescent on the (longitudinal) ribs; style 24-28 mm long, style dividing betweenthe middle and the apex of the anthers,style branches3.5-5 mm long. CapJUleand seedsnot seen

Specimensexamined. WESTERN AUSTRALIA: Vasse Highway, Busselton, I Sep. 1987, G.J. Keigherr9I50 (PERTH);old cemetery,Middleton Road,Albany, l4 Sep.1990, E. Croxford 6748 (PERTH);Moodong NatureReserve, 14 Sep.1992,G.J. Keighery 13519(PERTH);adjacentEastPerth Cemetery,Bronte St, Perth, 16 Sep. 1995,8.J. Lepschi &7.R. Lqlly l97l (PERTH).

Distribution- Native to the WesternCape Province,. In Westem Australia, B. nana is known from afew sitesin thePerth metropolitan area, as well asthe settlements ofBusselton and Albany. This speciesis probablymorewidespread in WestemAustralia than current collections would indicate. The first author has observeda naturalizedpopulation of this speciesat Kings Park, West Perth, but this is not supportedby a voucher specimen. It is likely that other naturalizedpopulations exist elsewherein the state,especially near settlementsin coastaland near-coastalareas in the south-west. (Figure1B)

Habitat. InWestemAtstraliaB. nanahasbeenrecordedgrowing in sandinAgonislexuosawoodlatd, Eucalyptusmqrginat( Banksia spp,woodland, and amongstvarious weeds in disturbedsites. Occurs on coastalsands in restioid fynbos (heathland)in South Africa.

Phenology. Flowering recordedduring August and Septemberin Westem Ausfialia.

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Breeding system. Within its natural ran ge,B. nanais pollinated by the large solitary bee Anthophorq diversipes Friese (Hymenoptera: Apidae) (Manning, unpubl'). Honeybees Apis melliftra L. (Hymenoptera:Apidae) havebeen observedgathering pollen from this speciesin Westem Australia (G.J.Keighery unpubl. obs.).

Notes. is superficially similar to B. angustifolia, and it is possible that it has been overlooked in Westem Australia as a result of confusion with the latter . Babiana nana has beenrefered to B. dlsticla in Western Atstralia in herb. (at PERTH) and in Husseyel al. ( 1997),but is not closelyrelated to that species. The earliestherbarium specimen ofB. nan.l datesflom 1987,but it is tikely that this specieswas naturalized in westem Australiasome years prior to this date. Although only Sparinglynaturalized in WestemAustralia at present,B. nanahas the potentialto becomea serious weedonsandy soils in thehigherrainfallareas ofthe south-westofthestate (G.J. Keighery pers. comm.). The presenceof a very large population spreadingthrough Banksia woodland in a natule Ieselve on the outskirts of Perth (see Keighery 13519) demonstates the threat this speciesposes to lemnant vegetation.

3, Sweet,Hort. Brit. ed. 1: 396 (1827).

BabianapulchraG.J.Lewis, J. S.African Bor Suppl 3:49 (1959)[asB. pulchra (Salisb ) G'J' Lewis].

2Babianoplicata auct. non (Thrnb.) Ker Gawl., Curtis's Bot Mag., 16: 1 576 ( 1802). See Cooke ( 1986a)and notesbelow.

Babianadisticha auct. non Ker GawL,Curtis's Bot. Mag.,16: t.626 (1803).

Babianastricta auct non (Ait.) Ker Gawl, Curtis'sBot Ma8., 16:t. 621 (1803).

Illustrations.Jeppe, Spring and Winter Flowering Bulbs of the Cape 104-105 (1989): Manning & Goldblatt,West Coast:S. African Wild Flower Guide 7: 68-69 (1996); Husseyetal , Westem Weeds 29 (1997) las B. stictal.

Perennialherb 15-60cmhigh. subgloboseto broadly ovoid, 10-30mmin diameter,tunics papery,decaying into coarseor fine fibres andextending up the baseof the stemfor 20-80 mm. Sten 5-20 --long. teaves distichous,spreading (fanwise) toerect, plicate, pubescent; sheathing base40- 200 mm long;lamina narrowly to very narowly elliptic, linear-elliptic,or rarely very narowly ovate, narrowly oblong-elliptic or narrowly oblong, 35-200 mn long, 4-22 mm wide, base attenuate- oblique, apexnarrowly acuteto acuminate.Inflorescencelonger than the leaves(often markedly so), 2-35-flowered,flowers twisted to facethe stemapex (so that the lowel(patterned) tepals are uppermost); scapesimple, l- or 2-branched,rarely much-branched,straight to occasionallyflexuose, pubescent' Brccts sheathing,herbaceous, reddish-brown, t narrowly ovateto very narowly ovateoI narrowly to very narrowly ovate-triangular(margins often i stlaight), or rarely narrowly oblong in lateral view, l5-30 mm long, pubescentto sericeous-pubescent,scarious apical portion glablous: apex acuminate to long-acuminate(frequently also lacerate), rarely narrowly acuteor aristate.Bracteoles free,loosely sheathing,herbaceous, brown to reddish-brown,t narrowly ovate in latelal view (margins often t shaight), 12-30 mm long pubescent,scarious apical portion and marginsglabrous; apex of individual bracieolesnarowly acute to acuminate. Perianth colour variable, usually purple to deep purple, bluish-purpleor violet, less often white (with pale blue flush abaxially), pinkish-purple or purplish- crimson, the tower three tepalseach with a dark red or blackish malk at the base(flequently with an ue -r{s '€ll€Jlsnv tuelsEeul (uoxEl pelsloJ 'elr e Jo Drloltsn8uD 'g ,(lqrssod)wuerqeq pelB^rllnJ;o sre,rog 3urlrsr,r.saaq,{euoq suode: oslu ( uuoc 'sred) pr seuol 1 q ('sqo Iqndun freq8roI I g) Erlullsnv uJe1se16ur sercedssrql uro{ uellod Auueql?8 'SuruuEI +I pe^rasqo uaeq e,ruq seeq,{euo11 ( lqndun ) (eEprdv :ereldouaru,{11)oseul sadrsn{p ,(t. poluu11od qlnos 'uatsKsSutpaatg onqdotpuy eaq,4Jelqosa8rrl eq1,{q sr sarcadssrgl'ecqJv u1 el!

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Nolex. The taxonfeated hereasB. ar?grsll/olia has previously beenrefened toB. plicatql= B, distichal (earlycollections in herb. aIPERTH),B. distichaor B. stricta (Green 1985,Cooke 1986a,Perry 1987). All threetaxa are similarin overall morphology,but may be distinguishedby variousfloral characters. ln B. distichathe pedanthtube is longer(18 25 mmasopposedtoT-18mmin B. angustifolia),and the stamensare more obviously arched. The flowers are also not twisted to face the stem apex (as in B. angustifolia), so the lower (marked) tepalsface away rather than towards the stem apex, and the flowers are often markedly zygomorphic. These featurescan be difficult to distinguish on dried material, and are best obseryed in live plants. may be distinguished from B. angustifoliaby the samecharacter offlower-odentation asdescribed above, as well asits somewhat broader,t saggitateanthers (anthers in B. angustifolia are narrowly oblong).

Nordenstam(1970) pointed out that the earliestname for thrs taxon,Acaste pulchra Salisb.,was not validly published and that the next available name is Babiana angusti;foliaSweet. The species teatedasBabianapulchraby Lew rs(195 9), the mostrecentmonographerofthegenus, is thuscorectly identified as B. angustifulia.

It shouldbe noted,however, that materialhere refered to B. angustifoliamay notrepresentthe 'true' wild variant of that species. Many speciesof Babiana are of horticultural interest (Bailey 1914, Harrison 1963, Bryan & Griffiths 1995), and the genushas been in cultivation in Australia since approximately the mid 19'r'Century (D.L. Jonespers. comm.). Naturalized babianasin Australia (including WesternAustralia) are presumablyderived from thesehorticultural introductions. In the caseof B. angustifolla,naturalized plants show considerablygreater variation in perianthcolour than do wild populationsofthis speciesin SouthAfrica, in which perianthcolour rangesfrom pale to deep bluish-purple or purple. The variability exhibited by naturalizedplants is probably the result of horticultural selection,possibly through hybddization with speciessuch asB. dlsticha or B. putpureo (Jacq.)Ker Gawl.,although this has not beenconfirmed.

ThenameB. s/rictahasbeen widely appliedto cultivatedbabianas formany years,bothin Australia (e.g.Pescott 1968, Hitchmough 1989) and overseas(e.g. Bailey 1914,Synge 1961, Harison 1963, Bryan1989). The conceptofB. aagustifoliahas var'redconsiderably between authors ofho icultural texts,being variously cited asa synonymof B. puLchra(e.g. Bryan 1989;cl Lewis 1959)regarded as a variety (var. angustifolia (Sweet) Baker) ofB. stricta (e.g. BatLley1914, Synge 1961) or treatedat specific rank (e.g. Bryan & Griffiths 1995). This suggestsa degreeof confusion with regard to the taxonomyofcultivated babianas,and clarification ofthe identity ofat leastthe more commonly grown taxa would be advantageous,particularly as this is the sourcefrom which the naturalisedmembers of the genusoriSinate.

Babiana angustfoLiais themost common and widespread sp ecies of BabianainW estemAustralia, and is also the most aggressivelyweedy. It is a major weed of heavy soils on the Swan CoastalPlain and the adjacent Darling Range area, often forming extensive monocultures to the detdment of indigenousherbaceous and bulbous plants. While presentlyonly naturalizedover a relatively small portion of the state,B. angustifolia has the potential to extend further into the more mesic coastal regions between Geraldton and Esperance; rainfall (along with soil type) appearsto be the main limiting factor affecting the distribution of this speciesin Westem Australia (G.J. Keighery pers. comm.).

The earliestcollection ofthis speciesfrom WesternAustalia seenby the presentauthors was made atGingin in 1925(R oad Board Secretarys.n.PERTH02022893). However, Ostenfeld( 1921)records "Babiana collecting plicata" on a roadside at Armadale h 1914 (Ostenfeld 166). This is almost tnI-I tE qDrylzssauraqulsuaplA 'e ',(u"rog 'ptquarso alsuo1 a3lp8uox )asppppaN aFtqopq uEd uErrEjrsnvrsatA or suoqnquruo) (tz6l) H t '€tt-€€t:EZl rasltoNlrlst otoq'ouolE)g pue Dgnorladr7:aeec?ppl u?rl{V qtnos uo seloN (0/61) g'lu€lsuaproN 69-19 :902 uounto^z puo srln watsts luold ?cl.+V ureqlnosq uretstrsuolt?ulllod,{U prrsoqord-8uol pezrtrrreds E pu€ sle^\olj peqnfSuol 'urtqptoD 'Suruuehl :pln8 uortBullod(eEprurtseu.raN:pratdr(\ sl.t1so.t!3uol snLlruittrolr!8aot4! eqJ, (t66t) d T Cf '6rl-l 'sr,{r.I :g lddns {uoloB uort4v q!1os{o lDurnol ouolqog snua} eqJ 16S6I) I D (,{euptrS:sseld sol!/rd qlnos ,raN Jo ,{trsro^run)Itl +II dd't IoA..set!tA qlnos^\oN Jo eroH,,Cpo) fC'uoprpH r4 eee3nprrl(€66I) vA'u^'\org T VJ'seurel ('Irvd euotcr^ :!rl?4snv uratse/llJo 'p,{on 'ppoc "C 'suesnoJ' ',&eqBra)' ',(essnH ,{Frros uoltelord tu?td) ,. spee/AuJelsel1\,, (166I) C S ? f U I D d ( Irotr ^\eN :uapl?5 Isrluelog > ol\ ^\eN) upa q18 ') 'ueiSurtoH ,{plror$ oqt Jo vlJ?qraHoql:l u?d unrol&qroH xepul, 10661) J.I lleul?g r[, HN uo€ruloH d ( poo^\8urd:IoN,O Suqr^) !{puvFez ^\aN puE €ll?rlsnv roJ sqlng uapre9,, (686I) I'q8nourqrllH '6 'uos^roH oN'sapes vun?J pue ?rolC usllvJlsnv.. ,{SolouluroJ,Io looqpusH V urntuounpullueld, (8861) IH ( qloN uotsreurled: oJ T uosurvH g'd) upe puz .. araqdsluoHureqtnos eqt roJ slvluuaradpuE sqlng Jo )tooqpu?H,,(€961) g d^'uosur?H ( quod :alnlln.lrBv Jo tuour[BdecuErlstsnv urelso/d upo puz ,,?llelsnv uelsey'AJo s]uvldrulncsv^ oq Jo snsueJ,,(9861) y'AI'uaoJD 'VJ 'raup&C ( qued |lelulrd tuaruue^oD) ,r'sllEtuoplcrooEllEl]snv ulnlgluDld ollereurnug,, (l€61) ( eurnoqlat{ :sserd eptuD 9ll*/89 dd'Z IoA,,tlolrr^ Jo rlotJ,,(spe) fI'etsl^ltug T DN'rlsle,4\iir.t c€oc?pll(i661) IB'uuoJ ( eprelopv iuorsr^rCSullulrd tueuure^oD uerle4s.nvqlnos) togl -Z8Lt dd't IoA,{slleJrsnvq]nos Jo srolC,t(spe) UH'ua:{trol 6' df'dossal rrl accceprrl(q986t) VO'e)looJ ( xraqutr) ierr^reS Surqsrrqnd '9t 'lo^ lueuue^oD uulejlsnv) 99FI dd .{plls4snv Jo erold,,Cpo) SV'o8moD rr1 erer?plrl (€9861) VO'e)tooo 'sqr$J!D CpullUod:ssord roqurl),,sqlng Jo lenu?'\I,,19661) ,,,1 T I'ur,tug 'usfig ( puxnrod;sserd roqurrl) I IoA ,, sqlng,, 1686t) f (llo^ ^{eN :uellrulrel^l) I lo^ .. onlln.ruoH Jo erpedotc,{)pr?puqs aql- ( 6l) Hl'^ol'Eg

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'Blota Perry, G.P. (198?). kidaceae. Inr Marchant, N.C. et aI of the Perth Region." Vol 2, pp 789-807' (West€m Aushalian Department of Agriculture: Perth ) "Bulbs Pescott, R.T.M. (1968). for Australian Cardens." (Nelson: Melboume.) philips, B.P. (1951). ,,Generaof south Aftican Flowering Plants." 2nd edn. Botanical survey Memoir No. 25. (Department of Agriculturci Pretoria.) "Colins Synge,P,M. (1961). Guide to Bulbs." (Collinsrt ndon.)