TRELITZI A ThisserieshasreplacedMemoirsoftheBotanicalSuweyofSouthAfricaandAnnalsof theKij’stenfroschBotanicGardenswhich SANBI inheritedfromitspredecessororganisa- tions. The plant genusSfrelitzia occurs naturallyin theeastern parts ofsouthernAfrica. It comprisesthreearborescentspecies, knownaswildbananas, andtwoacadescentspe- cies, known ascraneflowersorbird-of-paradiseflowers. ThelogooftheSouthAfrican National BiodiversityInstituteisbased onthestrikinginflorescenceofStrefifziareginae, a native ofthe Eastern Capeand KwaZulu-Natal that has become a garden favourite worldwide. Itsymbolisesthecommitmentofthe Institutetochampiontheexploration, conservation,sustainableuse,appreciationandenjoymentofSouthAfrica'sexception- allyrich biodiversityforall people. SCIENTIFICEDITOR: O.A. Leistner TECHNICALEDITOR: B.A. Momberg DESIGN &LAYOUT: E. Fouche COVER DESIGN: E. Fouche FRONTCOVER: Freeslafucata BACKCOVER: top: Freesiagrandi/Iorasubsp. grandi/bra middle: F. carymbosa bottom: F. verrucosa Citingthispublication MANNING,J.C &GOLDBLATT, P. 2010. Botanyand horticultureofthegenusFreesla (Iridaceae). Stnditzia27. SouthAfricanNationalBiodiversityInstitute, Pretoria. ISBN: 97S-I-9I9976-5S-7 ©Publishedby:SouthAfricanNationalBiodiversityInstitute. Obtainablefrom:SANBIBookshop. PrivateBagXlOl, Pretoria,0001 SouthAfrica.Tel.i +27 12843-5000. Fax: +27 [email protected] Website:www.sanbLorg Colourplates:BathopeleMarketing, P.O.Box72585, LynnwoodRidge,0040, Pretoria.Tel. (012)349-2951-Print- edby4lmages, PersequorTechnopark, BmmmeriaPretoria,P.O. Box34059. Glenstanna0010.Tel.012349-1113. TRELITZIA 27 Botany and horticulture ofthe genus Freesia •V (Iridaceae) by John C. Manning SouthAfricanNationalBiodiversityInstitute,PrivateBagX7,7735Claremont,CapeTown.UniversityofKwaZulu-Natal,Pieter- maritzburg.SchoolofBiologicalandConservationSciences.ResearchCentreforPlantGrowthandDevelopment,PrivateBag X101,Scottsville3209,SouthAfrica. & Peter Goldblatt B.A.KrukoffCuratorofAfricanBotany,MissouriBotanicalGarden,PO.Box299,St.Louis,Missouri63166,USA. with G.D. Duncan SouthAfricanNationalBiodiversityInstitute,PrivateBagX7,7735Claremont,CapeTown; F. Forest JodrellLaboratory,RoyalBotanicGardens,Kew,Richmond,Surrey,TW93DS,UnitedKingdom; R. Kaiser GivaudanSchweizAG,Uberlandstrasse138,CH-8600Dubendorf,Switzerland; Tatarenko I. JodrellLaboratory,RoyalBotanicGardens,Kew,Richmond,Surrey,TW93DS,UnitedKingdom. PaintingsbyAuriolBatten. LinedrawingsbyJohnC. Manning national biodiversity institute S A N B I Pretoria 2010 Several people helped materially by providing living material for il- lustration and we are very grateful to them for this: theyinclude Fanie Avenant from Victoria West, Fiona Barbour from Kimberley Anne Pa- tersonfrom Clanwilliam, Ted Oliverfrom Stellenbosch, members ofthe Kirstenboschbranch ofthe Botanical SocietyofSouthAfrica, andespe- ciallyCameronandRhodaMacMasterfromNapier, whopersonallycol- lectedanddeliveredfloweringandfruitingplantstousandtoAuriol.We alsothankElizabethParkerforherenthusiasmandforfacilitatingseveral collectingexpeditions, andRoseSmutsforhercompanyandhelpinthe field. Joop Doorduin, Freesia cultivar expert ofThe Netherlands, very kindly compiled the list of25 of the most popular cultivars. The elec- tronicmapswere kindlypreparedbyMichelle Smith and Les Powrie of the SouthAfricanNationalBiodiversityInstitute, andwearealsodeeply gratefultothe editorialstaffatthe Institutefortheirprofessionalism. Fieldworkforthis accountwassponsored inpartbyKatsumi Shimi- zu, who also provided notes onFreesia cultivation in Japan andwhose interest in this project was instrumental in its genesis. We are also ex- tremelygratefulto ElizabethParkerandtheParkerFamilyofElandsberg Nature Reserve for additional sponsorship towards the publication of thisvolume. Elizabethisastaunch and activesupporterofbotanical re- searchinSouthAfrica, withaparticularinterestinpromotingsystematic research into the flora. She participated actively in the expeditions to collect flowering plants for illustration, and her generous contributions expeditedthe appearance ofthisrevision. New c ecies and statuses in Strelitzia 27 (2010) . , Freesia grandiflora (Baker)Klattsubsp. occultaJ.C.Manning &Goldblatt, subsp. nov., 111 FFrreeeessiiaalperiacehtcloixniJi.KClMatatnsnuibnspg.&alGboladb(lGa.tLt.,Msepy..)noJv..,CM6a5nning &Goldblatt, stat. etcomb, nov., 70 Freesiasubgen. Viridibractea (Goldblatt)J.CManning &Goldblatt, stat. nov., 50 Freesiaviridis (Aiton) Goldblatt&J.CManning subsp. crispifolia (Goldblatt)J.CManning &Gold- blatt, comb, nov., 55 Auriol Batten We have great pleasure in dedicating this volume to Auriol Batten in recognition ofher unfailing talent and enthusiasm in this and other projectsthatwehavesharedwithher, notablytherevisionsof Gladiolus ‘ in southern Africa’ and of ‘Crocosmia and Chasmanthe. It has always been a great pleasure to workwithAuriol, and her clearvision and in- cisivemindhavebeenaninspiration. Shestartedworkonthisproject a yearbeforeherninetiethbirthdayandcompletedthepaintingstheyear — after aremarkable achievementbyaremarkable artist. The photograph depicts Auriol holding the Lifetime Achievement Award for Botanical Art, which she received at the Kirstenbosch Bien- nale in September2008. Auriol Ursula Batten is South Africa’s finest attheInauguralExhibitionofBotanicalArt, Kirst- living botanical artist. Born in Pietermaritzburg, enbosch Biennale, followed in 2008 bythe Life- KwaZulu-Natal Province, South Africa in 1918, time Achievement Award at the Kirstenbosch she graduated with a B.Sc degree from the lo- Biennale in recognition of her contribution to cal university in 1937, majoring in Botany and botanical artinSouthAfrica. Geography, and went on to pursue a career teaching at various schools in the Natal mid- Auriol has illustrated several botanical mon- lands. Herinterest inflowerpaintingstemsfrom ographs, including works on Dierama, Gladi- herschoolyears, whereshe receivedinstruction olus and most recently Crocosmia, and individ- fromtwodedicatedmistressesinbotanyandart, ualpaintingshave appearedinCurtis’sBotanical later studying art at the nearby Durban Techni- Magazine and Flowering Plants of Africa. Her cal College. She remained in the province until talents as an artist are matched byher scientific 1945 when she moved to East London, where insight and many of the plates in these works shelivestoday. owe theircompletenesstoherpersistence inac- quiringfruiting stages long after the initial work After settling down in East London she de- was done. Her workwas chosen for the exhibi- voted her spare time to painting and the result tionArtinScience atthe InternationalBotanical wasthepublicationin 1966ofWildflowersofthe Congress, Missouri in 1999 and is in the collec- Eastern Cape Province, a collaborative venture tions ofthe South African National Biodiversity betweenherselfand anotherlocal artist, Hertha Institute and ofthe Royal Botanical Garden, Ed- Bokelmann. This was the first wild flower guide inburgh. to the region and its genesis and production were due entirely to her creative and scientific Her interest in wild flowers extends far be- efforts. ThesumptuousFlowersofsouthernAfrica yond theirillustration. She is afoundermember followed in 1986, in which she illustrated 100 ofthe BorderWildflower Societyandstill serves species, again accompanied bytext written and ontheboardofvariousconservationbodies. For compiledentirelybyherself. Shewas awardeda these andhercontinued dedicationtobotanical Gold Medal bythe RHS that year and her inter- mattersingeneralshehasreceivedvariouscivic national reputation as a botanical artist was ce- decorationsandin 1994wasawardedanhonor- mented. In2000shewasawarded aGoldMedal arydoctoratebyRhodesUniversity. Contents Acknowledgements ii Newcombination,speciesandstatusesinStrelitzia27 (2010) ii AuriolBatten iii, iv Abstractandkeywords 1 Introduction 2 Materialsandmethods 4 Taxonomichistory 6 Morphologicalcharactersoftaxonomicsignificance 9 PhylogeneticrelationshipsbyF. Forest,I.Tatrenko,J. Manning&P. Goldblatt 22 Systematics 26 FreesiaEckl.exKlatt 26 Keytospecies 27 Subgen.Freesia 29 1.F. laxa(Thunb.) Goldblatt&J.C.Manning 29 la. subsp. laxa 31 lb.subsp.azureaGoldblatt&Hutchings 33 2.F. grandiflora (Baker)Klatt 33 2a.subsp. grandiflora Ill 2b.subsp.occultaJ.C.Manning&Goldblatt Ill 3.F andersoniaeL.Bolus 35 4.FspeciosaL.Bolus 37 5.F verrucosa (B.Vogel) Goldblatt&J.C.Manning 41 6.F. corymbosa (Burm.f.) N.E.Br. 43 7.F refracta(Jacq.)Klatt 45 8.F. occidentalisL.Bolus 49 Subgen. Viridibractea(Goldblatt)J.C.Manning&Goldblatt 50 Sect.AlataeJ.C.Manning&Goldblatt 51 9.F viridis (Aiton) Goldblatt&J.C.Manning 51 9a. subsp. viridis 55 9b.subsp.crispifolia (Goldblatt)J.C.Manning&Goldblatt 55 Sect. ViridibracteaGoldblatt 55 10.F.sparrmanii (Thunb.) N.E.Br. 55 1121..FF..fmuacragtianJa.taC.JM.aCn.nMianngni&ngGo&ldbGloaltdtblatt 5579 13.F caryophyllacea (Burm.f.) N.E.Br. 60 14.FpraecoxJ.C.Manning&Goldblatt 65 15.F leichtlinii Klatt 66 15a.subsp. leichtlinii 70 15b. subsp.alba(G.L.Mey.)J.C.Manning&Goldblatt 70 16.F.fergusoniaeL.Bolus 71 Additionalmaterialseen 76 Excludedspecies 79 FloralscentchemistryofFreesiaspeciesbyR. Kaiser 80 Pollinationbiology 86 Ecology 88 Biogeography 91 Speciationpatterns 91 Horticulturalhistory 92 CultivationandpropagationofFreesiaspeciesbyG.D. Duncan 96 References 104 — Appendix AnalyticalcompositionofFreesiascent 108 Addendum Ill Index 113 trelitzia 27 (2010) l Freesia Klatt (Iridaceae: Crocoideae) is charac- speciesin the subgenus comprise sect. Viridibrac- terizedbyitsregularlynettedcormtunics, sharply tea, definedbytwoderivedcharacters: apapillate deflexedorhorizontalflowerspike,deeplydivided stem and oblique seedswith swollen raphe. Spe- style branches, verrucose or papillate capsules, cies boundaries in sect. Viridibractea are particu- and subglobose seeds with a glossy, smooth or larlyproblematic, andtherecentdiscoveryofsev- lightlywrinkledtestalackingsecondaryornamen- eralnew, locallyendemicspecieshasnecessitated tation.Thegenuscomprises 16speciesdistributed a re-examination of the taxonomy of the group. through sub-Saharan Africa, with a marked cen- ThedevelopmentofaxillarycormelsinF.albaand tre ofdiversityin the winter rainfall region ofthe F. leichtlinii is apreviouslyoverlookedfeature that southwesternCape.Molecularandmorphological is diagnosticfor the two species, and populations evidencesuggestanewclassificationinwhichthe untilnowrecognizedatspecieslevelasF.alba,are tubular-floweredspeciespreviouslysegregated as distinguishable from F. leichtlinii onlyon the basis subgen. Anomatheca (Ker Gawl.) J.C.Manning & oftheir white rather than pale yellow or cream- Goldblatt do not comprise a monophyletic line- coloured flowers. They are accordingly reduced age but represent three independent origins for to subspecific rank as subsp. alba (G.L.Mey.) the gullet-flowered species. These data support J.C.Manning & Goldblatt. Subspecific rank is the recognition of two subgenera, based on the also re-introduced in F. viridis for inland popula- textureofthefloralbracts:subgen.Freesia (8spp.) tions with falcate, mostly crispulate leaves and isdefinedbyitssoft-textured, greenortranslucent shortlylanceolate tepals, which are distinguished floral bracts, oftenwith darkbrown tips; andsub- as subsp. crispifolia (Goldblatt) J.C.Manning & gen. Viridibractea (Goldblatt) J.C.Manning & Goldblatt from coastal populations with narrowly Goldblatt (8 spp.) is recognized by firm-textured, lanceolate leaves and attenuate tepals. The new green bracts with narrow translucent margins. species F. praecox J.C.Manning & Goldblatt is Floral chemistry provides additional novel data recognized for early-flowering populations with supportingthistopology: p-ionone and dihydro-p- narrowleavesfromthesouthernRiviersonderend ionone are characteristic ofallfragrantspecies in MountainsthatwerepreviouslyincludedinF.alba subgen. Viridibracteabutare absentfromsubgen. butwhich differfrom it in ecologyand in lacking Freesia. axillarycormels. Thisrevisioncoversmorphology, anatomy, molecular relationships, floral biology, In subgen. Freesia, plants of F. grandiflora taxonomy, horticultural history and cultivation. from Macuba, in north-central Mozambique, Each species is described and fully illustrated in with unusually short filaments and a consis- colour and black-and-white, with complete syn- tentlypinkperianth are recognised as subsp. oc- onomy, notes on ecology, and distribution maps. culta J.C.Manning & Goldblatt. Within subgen. Acompleteanalysisofthefloralscentchemistryof & Viridibractea, Freesia viridis (Aiton) Goldblatt 13 taxaispresented. J.C.Manningissegregated as the monotypicsect. Alatae, defined by several autapomorphies, no- Keywords: floral chemistry, Freesia Klatt, horti- tably a winged stem, greenish or brown flowers, culture, Iridaceae, newspecies, southernAfrica, anda 1-bandedpollenoperculum.Theremaining taxonomy 2 trelitzia 27 (2010) Freesia Klatt is one ofthe smaller ofthe Afri- The position of Freesia within the subfamily can genera ofIridaceae, alargefamilyofalmost Crocoideae has only recently become clear. The 2 000 species in ± 65 genera (Goldblatt &Man- divided style branches initially suggested a rela- ning 2008) and which includes popular garden tionship with the widespread African genus La- ornamentals such as Iris L., Crocus L., Gladi- peirousia Pourret, now placed in the subtribe olus L., Sparaxis Ker Gawl., Watsonia Mill, and, Watsonieae, andseveralofthenarrow-tubedspe- ofcourse, Freesia. The genus has a long history cieswereaccordinglyincludedbyBaker (1896) in ofcultivation in Europe, dating back to the late Lapeirousiasubgen.Anomatheca.Acloserelation- eighteenth century, and together with Gladiolus shipbetweenthesetwogenerahasnowbeendis- and Iris ranks as one ofthe horticulturally most counted:theaxialcormdevelopmentandsmooth important genera in the family Its colourful, seeds ofFreesia are fundamentally different from highly fragrant flowers have made it especially Lapeirousia, which has axillary corm develop- popular as a cut flower (Bryan 1995) and in re- ment, colliculate seeds, and unspecialized leaf hceanvteyreearasc,headnnouvaelrsa1l1e0s miniltlhieonUsntietmesd K(Fiungetdoalm. mGoalrdgbilnatatn&atoMmayn.niThnegs(e19a9n5a)totmoiscuaglgedsitfftehraetnFcreeselseida 2007). Astonishingly, the numerous cultivars might instead be allied to Tritonia Ker Gawl. and have been derived from repeated crosses be- CrocosmiaPlanch.,withwhichthepatternofcorm stpweeceinesjuasrtetawlomosspetciuenskannodwntheiinrcouflftsipvraitnigo.nWainldd dcoenvseilstoepnmte.ntS,ubsseeeqdusetnrtucatnuarleyasinsdolfeaDfaNnAatsoemqyueanr-e certainlynotin commercialnumbers. cesofseveralplastidregions(Goldblattetal.2005) showed thatFreesia is in fact immediatelyrelated Freesia, a member ofthe largelyAfrican sub- toCrocosmiaandDeviaGoldblatt&J.C.Manning, family Crocoideae, is recognized primarily by which have similar smooth seeds but notched, its sharply inclined to horizontal spike, deeply not deeplydivided style branches, and that these d(iGovliddbeldatsttyl&e brMaanncnhiesn,ga1n9d95v)e.rrOutchoesre cfaepastuulreess t(hGorledeblgaettn)erGaoldabrleatttog&etJh.eCr.Maallnineidngt,o wXheincohscaalpsao that characterize the genus are its mostly coni- showsthe unusual dividedstyle branchesbuthas unspecialized,angularseedswithasculpted,areo- lcaarlly(nreatthteerdcthoarnmstuunbigclso,boofstee)nccoormmpsosweidthofrfeignue-, late or foveate testa. Some species ofCrocosmia, as well as Deuia, have finelywrinkled and warty pale fibres. The leaves are anatomicallyunusual capsuleslikethoseofFreesia,providingindepend- in Crocoideae in lacking a submarginal vein or ent evidence for a close relationship between the sclerenchyma trace, with support provided in- three genera. ThefourgeneraFreesia, Crocosmia, stead by columnar, thick-walled epidermal cells Devia and Xenoscapa all share a similar, derived at the leafmargin (Rudall 1995). Other features leafmarginalanatomyandthederivedbasicchro- oafretthheegseunbugslotbhoasteasreeeddserwiivtehdsimnoottheh osrubsflaigmhitllyy mcoomsporimseinngumtrbieberxFr=ees1i1e,aaendwiatrheinnosuwbrfaemgialrydeCdroa-s wrinkled testa without visible cell outlines, and coideae (Goldblattetal. 2006). thebasic chromosome numberx = 11. The pol- len ofFreesia is typical ofsubfamily Crocoideae, Freesia is centred in the southwestern and with the striking exception of F. uiridis (Aiton) southern Cape in the southern African winter Goldblatt&J.C.Manning, whichhasa 1-banded rainfallregion,withtwospeciesextendingnorth- operculum ratherthantheusual2-bandedone. wards alongthe easternseaboardthroughtropi-