Bothalia A JOURNAL OF BOTANICAL RESEARCH May 2003 Vol. 33.1 TECHNICALPUBLICATIONS OFTHE NATIONALBOTANICALINSTITUTE, PRETORIA Obtainable from the National Botanical Institute, Private Bag XlOl, Pretoria 0001, Republic of SouthAfrica.Acatalogue ofall availablepublications will be issuedonrequest. BOTHALIA Bothalia is named in honourofGeneral Louis Botha, firstPremierandMinisterofAgricultureof the Union of South Africa. This house journal of the National Botanical Institute, Pretoria, is devoted to the furtherance ofbotanical science. The main fields covered are taxonomy, ecology, anatomy andcytology. Two parts ofthejournal and an index tocontents, authors andsubjects are published annually. Threebookletsofthecontents(a)toVols 1-20,(b)toVols21-25 and(c)toVols26-30,areavailable. STRELITZIA Aseriesofoccasionalpublicationson southernAfricanfloraandvegetation,replacingMemoirsof theBotanicalSurveyofSouthAfrica andAnnalsofKirstenboschBotanic Gardens. MEMOIRS OETHE BOTANICALSURVEYOESOUTHAERICA The memoirs are individual treatises usually ofan ecological nature, but sometimes dealing with taxonomyoreconomicbotany. Published: Nos 1-63(manyoutofprint).DiscontinuedafterNo.63. ANNALS OFKIRSTENBOSCHBOTANIC GARDENS A series devoted to the publication of monographs and major works on southern African flora. Published: Vols 14-19 (earlier volumes published as Supplementary volumes to the Journal of SouthAfrican Botany). DiscontinuedafterVol. 19. FLOWERING PLANTS OFAFRICA(FPA) ThisserialpresentscolourplatesofAfricanplantswithaccompanyingtext.Theplatesareprepared mainlybytheartists atthe National Botanical Institute. Manybotanical artistshavecontributedto the series, such as FayAnderson, PeterBally,Auriol Batten, Gillian Condy, Betty Connell, Stella Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700 plates), Claire Linder- Smith andEllaphieWard-Hilhorst. TheEditoris pleasedtoreceivelivingplants ofgeneral interest orofeconomic value forillustration. FromVol. 55, twenty plates arepublishedatirregularintervals. An indextoVols 1^9 is available. FLORAOFSOUTHERNAFRICA(ESA) A taxonomic treatise on the flora ofthe Republic of South Africa, Lesotho, Swaziland, Namibia and Botswana. The FSA containsdescriptionsoffamilies, genera, species, infraspecific taxa, keys to genera and species, synonymy, literature and limited specimen citations, as well as taxonomic andecological notes. Contributions to the FSA also appearinBothalia. PALAEOFLORAOFSOUTHERNAFRICA ApalaeofloraonapatterncomparabletothatoftheFloraofsouthernAfrica. Muchoftheinforma- tion is presented in the form oftables and photographic plates depicting fossil populations. Now available: Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, 1983, by J.M. & H.M. Anderson. Molteno Formation (Triassic) Vol. 2. Gymnosperms (excludingDicroidium), 1989, by J.M. & H.M.Anderson. Prodromus ofSouth African Megafloras. Devonian to Lower Cretaceous, 1985, by J.M. & H.M.Anderson.Obtainablefrom:A.A. BalkemaMarketing,Box317,Claremont7735, RSA. TowardsGondwanaAlive. Promotingbiodiversity andstemmingthe Sixth Extinction, 1999, byJ.M.Anderson(ed.) BOTHALIA AJOURNAL OF BOTANICAL RESEARCH Volume 33,1 Scientific Editor: G. Germishuizen Technical Editor: B.A. Momberg NASIONALE BOTANif^~S INISTITUUT PrivateBac- xin-' po-y.^n ',C, 2003 -G8- 3 NATIONAL 1 .Botanical INSTITUTE 2 CussoniaAvenue. Brummeria, Pretoria Private Bag XIOl, Pretoria 0001 ISSN 0006 8241 May 2003 CentenaryYear INCELEBRATION OF THECENTENARYOFTHE NATIONALHERBARIUM(PRE)* IN 2003AND ITSIMMEASURABLECONTRIBUTIONSTOBOTANYINSOUTHAFRICAANDFURTHERAFIELD EditorialBoard D.F. Cutler Royal Botanic Gardens, Kew, UK B.J. Huntley National Botanical Institute, CapeTown, RSA P.H. Raven Missouri Botanical Garden, StLouis, USA J.P. Rourke Compton Herbarium, NBI, CapeTown, RSA M.J.A, Werger University ofUtrecht, Utrecht,The Netherlands Acknowledgements to referees Acknowledgementstoreferees Archer, Mrs C. National Botanical Institute, Pretoria, RSA. Baijnath, Prof. H. University ofDurban-Westville, Durban, RSA. Bohs, DrL. University ofUtah, SaltLake City, USA. Brown, DrE. Royal Botanic Gardens, Sydney,Australia. Brummitt, DrR.K. Royal Botanic Gardens, Kew, UK. Burgoyne, Ms P. National Botanical Institute, Pretoria, RSA. Craven, Ms P. P.O.Box 399, Omaruru, Namibia. Crouch, DrN.R. National Botanical Institute, Durban, RSA. Edwards, DrT.J. University ofNatal, Pietermaritzburg, RSA. Hammer, S. SphaeroidInstitute, 2077 MonteVistaDrive, Vista, USA. Hilliard, DrO.M. Royal Botanic Garden, Edinburgh, Scotland, UK. Ihlenfeldt, Prof. H.D. Ostseestr. 40, D-24369Waabs, Germany. Kativu, DrS. University ofZimbabwe, Harare, Zimbabwe. Leistner, DrO.A. National Botanical Institute, Pretoria, RSA. Manning, DrJ.C. National Botanical Institute, CapeTown, RSA. Nicholas, DrA. University ofDurban-Westville, Durban, RSA. Nordenstam, Prof. R.B. NaturhistoriskaRiksmuseet, Stockholm, Sweden. Oliver, DrE.G.H. National Botanical Institute, CapeTown, RSA. Paterson-Jones, DrD. National Botanical Institute, CapeTown, RSA. Baton, DrA. Royal Botanic Gardens, Kew, UK. Relief, Ms E. National Botanical Institute, Pretoria, RSA. Smith, DrG.E. National Botanical Institute, Pretoria, RSA. Stedje, Prof. B. University ofOslo, Norway. Timberlake, DrJ.R. Biodiversity Eoundation forAfrica, Bulawayo, Zimbabwe. Van Wyk, Prof. A.E. University ofPretoria, RSA. Victor, Ms J.E. National Botanical Institute, Pretoria, RSA. Werger, Prof. M.J.A. University ofUtrecht,The Netherlands. .SeeBothalia28:271-297(1998). CONTENTS Volume 33,1 1. The genusSolanum (Solanaceae) in southernAfrica: subgenusLeptostemomim, the introduced sections Acanthophora and Ton’a. W.G. WELMAN 1 2. Studies in the genus Machairophyllwn (Mesembryanthemaceae), with notes on some related genera. H. KURZWEILandP. CHESSELET 19 3. Anew serotinous species ofCliffortia L. (Rosaceae) from Northern Cape, South Africa and section emended.A.C. EELLINGHAM 41 4. Arevision ofLedebouria (Hyacinthaceae) in SouthAfrica. 3. The reinstatement ofL. ensifolia, L. gol- pinii andL. sandersonii. T.J. EDWARDS and S. VENTER 49 5. An annotatedchecklistofthe pteridophyte floraofSwaziland. J.P. ROUX 53 6. Taxonomy ofthe genus (Thymelaeaceae). C.L. BREDENKAMPandA.E. VANWYK .... 59 7. Studiesinthe liverwortfamilyAneuraceae (Metzgeriales)from southernAfrica. 5. Riccanliaamazonica. S.M. PEROLD 99 8. Notes onAfrican plants: Apocynaceae. Anew subspecies ofBrachystelma from Eastern Cape, SouthAfrica(Asclepia- doideae-Ceropegieae).A.P DOLD and PV BRUYNS 105 Asteraceae. Anew species ofPhilyropliylliim (Gnaphalieae) fromNamibia. PPJ. HERMAN .... 118 Hyacinthaceae.Anew species andnewcombinations inDrimia (Urgineoideae). J.C. MANNING andP. GOLDBLATT 109 Hyacinthaceae. Anew species ofOrnithogaliim from Eastern Cape, South Africa. A.P. DOLD HAMMER and S.A. 112 Hyacinthaceae and Crassulaceae. Two new cremnophilous taxa from semi-arid regions in South Africa. E.J. VANJAARSVELD andA.E. VANWYK 115 Lamiaceae. Tetradenia kaokoensis, anew species from Kaokoland, Namibia. E.J. VANJAARS- VELD andA.E. VANWYK 107 Lamiaceae. The correct name forSalvia thermara. E.J. VANJAARSVELD 112 9. AcomparisonofMopaneveldvegetationinSouthAfrica.NamibiaandZimbabwe. F. SIEBERT,G.J. BRE- DENKAMPand S.J. SIEBERT 121 10. Miscellaneous notes: Hyacinthaceae. Chromosome studies onAfrican plants. 19. New chromosome counts for three taxa. P.D. LEBATHA, J.J. SPIES and M.H. BUYS 135 11. Bookreviews 139 New name, species, subspecies and combinations in Bothalia 33,1 (2003) Albucatherniarum VanJaarsv., sp. nov., 116 BrachystelmafranksiaeN.E.Br. subsp. grandiflorumA.PDold & Bniyns, subsp. nov., 105 CliffortiadichotomaFellingham, sp. nov.,41 Crassulafoveata VanJaarsv., sp. nov., 116 Drimia barkerae Oberni. exJ.C.Manning <& Goldblatt, sp. nov., 109 Drimia ciliata (Lf.)J.C.Manning & Goldblatt, comb, nov.. Ill DrimiahesperanthaJ.C.Manning & Goldblatt, nom. nov.. Ill Drimia kniphofloides {Baker)J.C.Manning & Goldblatt, comb, nov., Ill Drimia nana (Snijinan)J.C.Manning & Goldblatt, comb, nov.. Ill Drimiasaniensis (Hilliard & B.L.Burtt)J.C.Manning & Goldblatt, comb, nov.. Ill Ledebouria ensifolia (Eckl.) S.Venter& T.J. Edwards, comb, nov.,49 Ledebouria galpinii (Baker) S.Venter& T.J. Edwards, comb, nov., 50 Ledebouriasandersonii (Baker) S.Venter& T.J. Edwards, comb, nov., 50 Ornithogalum perduransA.P.DoW& S.A.Hammer, sp. nov., 112 PasserinaesterhuyseniaeBredenkamp &A.E.van Wyk, sp. nov., 75 Philyrophyllum brandbergenseP.P.J.Herman, sp. nov., 118 Tetradenia kaokoensis VanJaarsv. &A.E.van Wyk, sp. nov., 107 iv Bothalia33.1: 1-18(2003) The genus Solanum (Solanaceae) in southern Africa: subgenus Leptostemonum, the introduced sections Acanthophora and Torva W.G. WELMAN* Keywords:AcanthophoraDunal,ethnobotany,Leptostemonum(Dunal)Bitter,morphology,Solanaceae,SolanumL.,southernAfrica,taxonomy, Ton'aNees ABSTRACT InthegenusSolanumL.(Solanaceae),subgenusLeptostemonum(DunalIBitter,sectionAcanthophoraDunalhasfour representativesintheFloraofsouthernAfricaregion(SouthAfrica,Lesotho,Swaziland,Namibia,Botswana),namelyS. aculeatissimumJacq.,S. Allioni.S.mammosumL.(cultivatedonly)andS. viarumDunal.Section TonaNees hastworepresentativesinsouthernAfrica,namelyS. chrysotrichumSchltdl. (S. hispidumauctt. nonPers.)andS. tonmni Sw.;botharenaturalizedweeds.Solanumcapsicoides,S. viarumandS. ton'um havenotbeenlistedbeforeforsouthern Africa.AllareintroducedspeciesnativetotheNewWorld.Descriptions,discussions,illustrationsanddistributionmapsof thenaturalizedspeciesarepresented,aswellaskeystothespeciesofbothsections. CONTENTS Detailed descriptions and discussions are given forboth sections and all six species, together with keys, illustra- Abstract 1 tions and distribution maps. S. mammosum, being culti- Introduction 1 vatedonly, has no distribution map. Descriptions anddiscussions 1 SectionAcanthophora Dunal 1 Solanum aculeatissimum Jacq 3 DESCRIPTIONSANDDISCUSSIONS S. capsicoidesAllioni 6 S. mammosum L 8 Section Acanthophora Dunal, Histoire naturelle, S. viarum Dunal 10 medicale et economique des Solanum: 131, 218 (1813). Section Torx’a Nees 11 Lectotype species: S. mammosum L. (D’Arcy 1972). Solanum chrysotrichum Schltdl 12 S. tonnim Sw 13 Section Aculeata G.Don subsection Acanthophora Specimensexamined 15 (Dunal) G.Don: 434 (1838). Acknowledgements 16 References 16 Psilocarpa, grad, ambig., Dunal: 216 (1852). Section Simplicipilum Bitter: 147 (1923). Lectotype INTRODUCTION species: S. aculeatissimum Jacq. (Seithe 1962). In the genus Solanum L., the prickly subgenusLepto- Description (basedon Nee 1991) stemonum (Dunal)Bitterisrepresentedbyeightsections insouthernAfrica.Twosectionscontainonlyindigenous Small shrubs, 0.3-3.0mtall, sprawling ormore often taxa, one section contains mainly indigenous taxa, and erect and few-branched, annuals or short-lived perenni- the remaining sections have only introduced species. als, heavily armedwith straightacicular(needle-shaped) Section Acanthophora Dunal is represented by four prickles, in some species stems with additional robust species in southern Africa. All are originally from the recurved prickles from enlarged compressed bases. Americas, namely the rather common and widespread Stems usually glandular-puberulent, often also eglandu- weedS. aculeatissimum, the relatively rare weedS. cap- lar-pilosewithfew-celled,simplehairs.Leavespetiolate, sicoides,thecultivatedornamentalS. mammosum,andS. alternate or usually geminate, with one about twice the viarum which is becoming a fairly common weed in size of the other; blades usually ovate to orbicular and KwaZulu-Natal and neighbouring areas. S. capsicoides with 2-4 pairs of lobes, or repand, coarsely and irregu- and S. viarum have not been listed before for southern larly toothed, membranous, spiny along main veins Africa (Welman 1993). Section Totra Nees is represent- above and below, usually glandular-puberulent above edby two species in southernAfrica; both are originally and usually also eglandular-pilose with simple few- fromtheAmericas.S. chrysotrichum issofarnaturalized celled hairs, these sometimes with vestigial rays at base, only in Eastern Cape, while 5. tonnun is confined to a below with simple hairs and sessile few-rayed stellate fairly small area north of Durban, KwaZulu-Natal. hairs(theselackinginS. capsicoides).Inflorescenceslat- eral, simpleorfew-branched, sessileorshortly peduncu- late,remotefromleaves, axisoften spiny; flowersfewto 0*0N0a1tiPorneatloriHae.rbarium.NationalBotanicalInstitute,PrivateBagXlOl, many, fewerthan halffertile, often only lowest fertile in E-mail:wgw(s>nbipre.nbi.ac.za large-fruited species. Calyx seldom much accrescent; MS.received:2001-06-28. lobestriangulartolinear. Corollawhite,yellowish,green- 2 Bothalia33,l (2003) ish orblue-puiple, glandular and pilose on outside, stel- only present onjuvenile plants or leaves. During evolu- late, without interstitial pleating. Ovary glabrous or tion the basal side rays ofthe stellate hairs have appar- minutely stipitate-glandular, puberulent in few species ently disappeared. Single-celled simple hairs, often with simple hairs that drop offearly. Fruit usually glo- gland-tipped,may alsooccurontheuppersurfaceofthe bose, small to large, strongly dark green and pale green leafas well as elsewhereontheplant(Nee 1979). marbled when young, yellowish, orange or reddish at maturity, glabrous at maturity, exocarp ± tough, meso- Diagnosticcharacters carpjuicy, in some species becoming dry at full maturi- ty, placentae simply columnar, divided or variously AccordingtoWhalen(1984),theS. mammosumGroup intruded, the fruit becoming effectively or completely ofspecies is distinguished by an essentially herbaceous unilocular at maturity. Seeds numerous, subreniform to habit, difoliate-geminate sympodia, vestitureofpredom- orbicular, seed coat closely investing endosperm or inantly simple hairs, variously lobed or dentate, prickly extendingaroundperipheryofseedintoaflattenedwing. leaves, lateral, often simple inflorescences, stelliform corollas anddryish, glabrousberries. Distribution andecology Nee (1979) stated that section Acanthophora, as a memberofthepricklysubgenusLeptostemonum(Dunal) According to Whalen (1984), section Acanthophora occurs throughout the American tropics except in Bitter, deserves its name; all the species are heavily armed with sharp, slender prickles and some species AAnmdaezsonainadninBrsaozuitlh;eaistteirsnmBorasztil.diAvbeorsuet2in0stpheecineosrtghreorwn have stouter, broad-based prickles as well. He defined the section by the combination ofthe upperleafsurface in open savanna and disturbed sites. Several species are established in the OldWorldtropics. bearing only simple hairs, glabrous fruit and thenon- to only slightly accrescentcalyx. Nee (1979) noted that this section, in common with This section can be divided into two subsections almost all groups ofSolanum, has great weedy tenden- based on the seedmorphology (Nee 1979). One subsec- cies; the species are found in areas ofhigh light intensi- tion, with seeds that do not differ much from those of ties and also where the natural vegetation has been dis- almostallotherSolanum species,includesS. aculeatissi- turbed. Under natural circumstances, landslides, eroded mum,S. mammosumandS. viarum.Theothersubsection stream banks and forest openings provide suitable habi- is characterized by seeds surrounded by a broad, flat- tats. Man’s agricultural and industrial practices are still tened margin or wing, a character unique in Solanum. expanding the ranges ofnumerous species ofSolanum. The succulent berries ofmany species are popular with TS.hiscapssuibcsoeicdteiso.n iTshperoebvaoblluytimoonnaorpyhysliegtniifcicaanndceinocfludtehse birds andensure seeddispersal. winged seed is not clear; it could aid in wind or water dispersal. Taxonomy Sexformsandbranching Dunal (1813) appropriately named this group of prickly speciessectionAcanthophora (thorn-bearing). In Whalen & Costich (1986) described section Acan- 1923, Bitter referred to the characteristic hairs of this thophoraasweaklytostronglyandromonoecious;having section in the name Simplicipilum. However, Dunal’s both perfect and male flowers, but no female flowers. name forthis section musttakeprecedence. Species of this section have some short-styled flowers, and are self-compatible. The fruits are small to large; it In 1972, D’Arcy elevated subsection Lasiocarpum was found that andromonoecy in section Acanthophora Dunal ofsection Acanthophora to section Lasiocarpum is most strongly developedin large-fruited species. (Dunal) D’Arcy, with S. lasiocarpum Dunal as type species. Section Acanthophora can be distinguished Child (1979) and Child & Lester (1991) studied the fromsectionLasiocarpumbythelackofstellatehairson branching patterns in the Solanaceae. All sections ofthe theovaries, the moregeneral coverage with multi-celled subgenus Leptostemonum have 2- or3-foliate sympodial simplehairs and the less substantial habit (Jaeger 1985). unitsoftenwiththesubtendingleafpairedwiththesmall- Section Lasiocarpum is indigenous to South America erfirstleafofthedaughtershoot(anisophyllousgeminate and no representatives have so far been recorded from sympodia). Section Acanthophora has mostly simple southern Africa, though a few species are naturalized few-flowered inflorescences with only the basalmost and/orcultivated in EastAfrica; see Jaeger(1985). flowerand/orthe few proximal flowers ofthe subsidiary inflorescencehermaphrodite;thedistalflowersareusual- Whalen (1984)placedS. aculeatissimum, S. capsicoi- ly smaller, brachystylous or short-styled (andromonoe- des, S. mammosum and S. viarum in his 5. mammosum cious; Symon 1979) and are often inserted with a spatial Group. gap from the hermaphrodite flowers. Species with prick- ly calyces in the hermaphrodite flowers usually have unarmed, functionally male, brachystylousflowers. Microscopestudies Section Acanthophora has distinctive simple hairs Note that are long, multicelled, uni,seriate, shiny and translu- cent.Thesesimplehairsareidentical with thecentral ray Nee (1991) noted that this section contains a number of the stellate hairs on the same plant; the latter often ofpromising candidates forscreeningforalkaloids. . Bothalia33.1 (2003) 3 Keyto species ofsectionAcanthophora in southern diam., brownish, moderately flattened, seedcoat closely Africa (fromJaeger 1985) investing endosperm, with margin sometimes thickened la Seeds 4-6 mm diam., prominently winged; branches and butnot wing-like. Figure 1 leavesalmostglabroustosparselyhairy;ripefruitVer- million(sometimescultivated) S.capsicoides Distribution lbSeeds2-5 mmdiam.,notwinged;branchesandlowerleaf surfacehairy;ripefruityellow: 2aStemswithacicularpricklesonly;brancheswithscattered Whalen (1984) gave the distribution ofthis species as simplehairs S.acideatissimiim southea.stem Brazil (natural) and Central Africa (natural- 2b Stems prickly with both compressed and recurved and ized). Jaeger (1985) stated: The wide distribution ofthis long,straightacicularprickles;brancheshirsute with speciesinAfrica,anditsscarcityorevenabsencefromthe 3aHaidresnsoen,ssitmepmleofhaviarrsi:able lengths, 2^ mm long; calyx New World, together suggest that S. aculeatissimum is not prickly; corolla violet; fruit globose, usually nativetoAfrica.ThisgivessectionAcanthophoraaninex- with an apical nipple and several basal protuber- plicablydisjunctdistribution.However,S. aculeatissimum 3bHairsaprnoicncekssl;tye;srenceoodrfsoul5lnaimfwomhrimtdeil;aenmfg.rtuhi(stc,uglultpoibvtoaostee5d;mosnmeleydl)son±g;S.2cmamlaymmxmosum giissrcomowonsrseiadsleiarkeewldeytehtdaotobtfersaaednecoirnnodtuartoredsyubcveteigtoewtneaettonioAtnfh,reiiNcnade;iwcaanWtdionrwglhtdehaantniidtt diam S.viarum Africa have been established for over 400 years (Roe 1979), an early introduction ofthis species would allow 1.SolanumaculeatissimumJacq.,Collectaneaaus- tmiumsetfdoerteitrsgsruabzsienqgueanntddciosnpterrisbault.eIttsohiitsdesouuccselsys.s’pinyhabit triaca ad botanicum 1: 100 (1787a); Jacq.: t. 41 (1787b); GB(Rib1eti9ltl8iee8er:)f:;11&N145e8e(P(:1.1P992.7J269.35)H);;e(rJ1Ham9ee9aig1nne):er;:;B436u723k824e((n1(1y919a896593&)7););;CBPauerSakarhessancewoy:::a15&72133(H4(a1,1l99l7:92853)8)9;4; mrienuggJimaaocenshgseieeerimvn&esdwHteaoesptvheeparrevnyre,w(bie1eda9ees8nt6de)iirnsnctAorfnairfbniuidctriamsoenfodouirtnthhateahrtlenoSAn.Agfafrcrtouiilmcmeaoea.,nttihsNaasenviee-- (2000).Type:cultivatedplantatVienna, ‘Patriainzonator- (1991) reported that S. aculeatissimum occurs naturally ridaest’,Jacquins.n. (W). mainly from Rio Grande do Sul, Parana and Santa S. reflexionSchrank: t. 81 (1822).Type:cultivatedplantinHortus Catarina states in Brazil. He also reported that this Monacensis(Munich),vonSchranks.n.(M). species has long been grown in European botanical gar- denswhere itis very uniform andmay representasingle NunkS.lokwh.asJ.iDa.nHuomokCe.Br.&ClaTrTkhe:om2s3o4n(11488(3)C.ALT)y.pe: India, Khasia Mts. introductionfromSouthAfricanpopulations, whichthey strongly resemble. S. cavaleriei Leveille & Vaniot: 207 (1908). Type: from China. J.Cavalerie2722(E). Because of confusion with S. capsicoides and other heavily armed species of Solanum, distribution records Annual, sparsely branched herb or undershrub up to 1.5 m high, pubescence highly variable, dense to sparse oabfleS.. IancusloeuatthiesrnsiAmfurmician,tthhieslsipteecriateusreisafraeirolfytcenomnomtonreliin- (plant rarely nearly glabrous). Stef7is tinged purple and Limpopo [Northern Province], Mpumalanga, KwaZulu- green, rarely densely pubescent, with simple, umnimseriate, Natal, Eastern Cape and rare in the Free State, Lesotho multicellularhairs ofunequal lengths, up to 4 long, andWesternCape, withoneisolatedrecordinthe North- heavily armed with many slender, acicular, mstmraight, West almost on the borderwith Gauteng (Figure 2). It is swmipmtr,heabbdaaissneeg0ot.rr5usnlmciagmthetlboyrrorasedlf.ilgehLxteeladyvpcerosircdokavltaeets,e,ulpoubptaottio1o2n16v0arxialb1ol8ne0g, ega.lrgso.owinsnoambteottthaienmieBcsoatlacnugilatcriadvleantSset;datiaFosonr,abDecusurribNoaHsni3t4yin6i4n149t4h4(e.NseH)arewaass, usually doubly or dentately lobed to 'A width of leaf, with 3 or4pairsofmajoracute, triangularlobes, middle lobe up to 55 mm long, dark green when fresh, usually Ecology drying dull dark green, with simple pilose hairs on both surfacesofleaves,rarelywithstellatehairsonunderside, Nee (1991)recordedthatS. aculeatissimum isaweedy mm prickles on midrib and petiole slender, ± 15 long, shrubthatgrowsindisturbedorpasturedforest,ingrazing base ± 2 mm broad, prickles on primary lateral veins fieldsandalongroadsides,from400-1 200maltitude. Tn mm slender, ± 7 long, purple near base, upper part yel- sub-SaharanAfricaitisfoundatforestedgesandinsavan- low, the base 0.8 mm broad; petiole 4-80 mm long. nas,mostlyintheeasternuplandsofthegreatlakesregion, Inflorescences simple, 3-5(6)-flowered; peduncles up to butscatteredthroughtherainforestandintoSouthAfrica, 0.5 mm long; pedicels± 10mm long. Calyx: lobes often from 1 000-2400 m, rarely descending almost to sea unequal, elongating immediately after anthesis and con- level.’Gbile(1979)describedthisasahighlandspeciesin cealing immature fruits with subcaudate lobe tips, Nigeria that grows mostly on rough ground and amongst pubescent, becoming prickly in fruit. Corolla 10-15 x rocks.AccordingtoBukenya&Carasco(1995),S. aculea- 25-30 mm, stellate, white, greenish or yellow-green, tissimum generally grows in forest clearings in Uganda; sometimes with a purple tint, rarely with purplish hairs. accordingtoBukenya& Hall (1988),itisararespeciesin Stamens:filamentsgreentoyellow;anthersyellow.Style secondaryforestin western Ghana. white; pistil pale green. Fruit nodding, globose, 15-30 mm diam., smooth, pale green with medium green In southern Africa, S. aculeatissimum grows in forest stripes or white with green reticulation when young, margins and clearings, along forest paths, and among dirty yellow or brownish when ripe. Seeds 2-4 mm undergrowthinclosednaturalforest,butalsoinFinnsand 4 Bothalia33,1 (2003) — FIGURE1. A,Solcmumaculeatissi- mum, Acocks 10787, fruit Nienaber9(PRE);B,S. tor- vum, Henderson 737(PRE). Artist:G.Condy. Eucalyptus plantations. It prefers damp, shady places February, similarly in East Africa, but throughout the undershrubsandtrees.Itisalsofoundalongroadsidesand yearin theDemocratic Republic oftheCongo. In south- in other disturbed areas, as a weed in gardens, ploughed fields as well as grassland. Wells et al. (1986) listed S. aculeatissimurn as a ruderal, silvicultural and pastoral weed of the temperate and subtropical summer rainfall areas in southern Africa. It is found on gentle and steep mountainandhillslopes,ingorgesandvalleys,butalsoon flats; it can grow on moist humus-rich and red-brown loamysoil andalsoon well-drainedsandyorstony soil. It is common in mountainous areas and is recorded from about sealevel to± 2200m altitude, with arainfall ofup mm to2000 annually.Acocks(1988)listedS. aculeatissi- miiin as one ofthe generally occurring smaller plants of the forest floorand margin in his Northeastern Mountain SourveldwhichisoneoftheInlandTropicalForestTypes. He also listed it as one of the smaller plants of general occurrence in the Highland .Sourveld which is one ofthe Temperate andTransitional Forestand ScrubTypes. Nee (1991) stated that in Brazil, its natural distribu- FIGURE 2.—Distribution of Solanum acideatisshmim in the FSA tion area, flowering is confined from October to region.