SystematicBotany (2004),29(1): pp. 177– 187 q Copyright 2004by the AmericanSociety of PlantTaxonomists AChloroplast DNAPhylogeny of Solanum Section Lasiocarpa LYNN BOHS DepartmentofBiology ,University ofUtah, Salt Lake City,Utah 84112 ([email protected]) CommunicatingEditor: J amesF .Smith ABSTRACT. Solanum section Lasiocarpa includes about adozen species witha center ofdiversity in the New Worldtropics. Solanum lasiocarpum and S. repandum (sometimesconsidered tobe conspecic as S. ferox)have an OldW orlddistribution in Asia and the Pacic Islands.Several species inthis section produce edible fruits,and two, the lulo ornaranjilla ( S.quitoense ) and the cocona ( S.sessiliorum )are cultivated commercially.Phylogene tic relationships in Solanum section Lasiocarpa were investigated usingsequence data fromthe chloroplast trnT-trnL spacer, the trnL-trnF spacer, and the trnL gene, including the trnL intron.Sampling included 24 accessions fromsection Lasiocarpa and 14 accesssionsofother Solanum species as outgroups.Allspecies considered tobelong to section Lasiocarpa bypreviou sauthorsw ere examined withthe exception of the recentlydescribed S. atheniae.Solanumrobustum and S. stagnale,sometimesconsidered to belongto section Lasiocarpa, are excluded fromthe group on the basis ofthe trn data. The remainingspecies inthe section forma monophyletic group,with threew ell-supportedclades withinit: S. hirtum, S.pectinatum -sessiliorum -stramonifolium ,and the remainderof the species inthe section. Sequencesof S.lasiocarpum and S. repandum are extremelysimilar, and these twoAsian taxa cluster withthe New World S. candidum and S.pseudolulo on the trn trees. Solanum section Lasiocarpa (Dunal)D’ Arcy compris- group ofthe genus Solanum,usually recognized as So- es approximately adozen species ofperennial shrubs lanum subgenus Leptostemonum .Previous authors such or small trees with acenter ofdistribution in north- asDunal (1852),Seithe (1962),Danert (1970),D’ Arcy western SouthAmerica. Morphologicalcharacters that (1972),and Whalen (1984)have regarded subgenus dene the section include difoliate sympodial units, Leptostemonum asa naturalgroup based on the shared large repand leaves, unbranchedinorescences ,stel- presence in most species ofspines, stellate hairs, and latecorollas, an dfruits covered with stellate hairs with tapered anthers. Molecularph ylogeneticstudies based reduced lateral rays (Whalen et al.1981). The section on chloroplast DNArestriction sites (Olmstead and wasmonograph ed byWhalen et al.(1981), who rec- Palmer, 1997)and nuclear and chloroplast sequence ognized 13species. Eleven are native tothe northern data(Bohs and Olmstead 1997,1999, 2001; Bohs, in Andes ofV enezuela,Colombia, Ecuador, and Peru, and press) indicate that Solanum species thatbear spines as three haveranges thatextend into Central America ( S. well asstellate hairs comprise amonophyletic group, candidum, S. hirtum)or northeastern SouthAm erica termed the Leptostemonum clade byBohs (in press). through the Guianasinto northern Brazil ( S.stramoni- Solanum wendlandii ,arepresentative of Solanum section folium).Several species in the section produceedible Aculeigerum Seithe,fallsoutside the cladecom prised of fruits and two, S.quitoense (the luloor naranjilla) and the other spiny Solanum taxa(Bohs and Olmstead 1997, S.sessiliorum (the cocona)are economicallyimportant 1999,2001; Bohs, in press). Solanum section Aculeigerum fruitcrops in Latin America (Heiser 1969,1985a). So- includes six species thatbear spines butlack stellate lanum quitoense has been introduced toPanama, Costa hairs. In this paper, Solanum subgenus Leptostemonum Rica,and Guatemalaand is now naturalized in Central is used in the traditional sense torefer toall taxa of America. Solanum lasiocarpum and S. repandum are the genus thatbear spines. The term Leptostemonum found in Asia and the Pacic Islands. Although treated clade is used in accordancewith Bohs (in press) to asseparate taxa by Whalen et al.(1981), H eiser (1996) refer tothe monophyletic group ofspin y Solanum taxa considered them conspecic under the name S. ferox. exclusive of Solanum section Aculeigerum . Dunal (1852),M orton (1976),and Hunziker (2001) Molecularstudies based on chloroplast DNArestric- included the SouthAm erican S. robustum in section La- tion sites and chloroplast ndhF sequence datausing a siocarpa,butWhalen et al.(1981) exclude dit from the broadrange ofsampling from Solanum indicate that section due todifferences in branching pattern, leaf section Lasiocarpa may bea relativelybasallineage shape,and fruittrichom es. Subsequent toWhalen et within the Leptostemonum clade and thatit may be al.’s (1981)treatment, S. stagnale wasremoved from the sister to Solanum section Acanthophora Dunal (Olmstead section and placedin the S.polytrichum group within and Palmer 1997;Bohs, in press). Whereas these broad Solanum subgenus Leptostemonum (Dunal)Bitter scalestudies sampled only one totw ospecies from the (Whalen 1984;Child 1998;N ee 1999).Symon (1985) section, species-level relationships in section Lasiocarpa described S. atheniae from NewGuineaand postulated have been the subjectof n umerous investigations using thatit belonged tosection Lasiocarpa. morphologicaldata, crossing studies, isozyme electro- Solanum section Lasiocarpa belongs tothe spinysub- phoresis, karyotype analyses, and cpDNArestriction 177 178 SYSTEMATIC BOTANY [Volume 29 TABLE 1. Sources of Solanum DNAaccessions used inthis study. Seeds, leaves, orD NAextracts provided by 1 L. Bohs, University ofUtah, Salt Lake City,UT. 2 R. G.Olmstead,U niversityof W ashington, Seattle,WA. 3 A. Bruneau, McGill University,Montreal, Canada. 4 C. B. Heiser, Jr., Indiana University,Bloomington,IN. 5 J.Miller, Amherst College ,Amherst,MA. a Forfurther collection and voucher data see Appendix inWhalen et al. (1981). BIRMsamples bear the seed accession numberof the Universityof Birmingham Solanacea e collection. Nijmegenaccession numbersrefer to the Solanaceaecollection atthe Universityof Nijmegen, The Netherlands. Solanum section Lasiocarpa:S. candidum Lindl.3—Stoutamires.n. (IND) from Heiser S249a,Mexico: Veracruz (AY266250). S. candidum Lindl.1—Bohs 2898 (UT),Costa Rica: La Cangreja (AY266237). S. felinum Whalen4—Benitez de Rojas 8915 (IND), Venezuela: Colonia Tovar (AY266252). S. hirtum Vahl3—Whalen 730 (QCA),Ecuador (AY266254). S. hirtum Vahl3—Jones s.n. (IND) from Heiser S404a,Costa Rica: Guanacaste (AY266253). S. hyporhodium A.Braun &Bouche´3—Whalen 717 (BH), Vene- zuela: Sucre (AY266238). S. hyporhodium A.Braun &Bouche´4—Carreno Espinosa 8214 (IND),V enezuela: Sucre (AY266255). S. lasiocarpum Dunal4—Ansyar 9605 (IND),Indonesia: Pandang (AY266256). S. pectinatum Dunal4—Peeke 8512 (IND), Ec- uador: Limoncocha (AY266227). S. pectinatum Dunal1—Bohs 2899 (UT),Bolivia: Santa Cruz (AY266230). S. pseudolulo Hei- ser3—Plowman etal. 4276 (GH)a,Colombia: Meta, Sierrade la Macarena (AY266258). S. pseudolulo Heiser5—BohsDNA ex- tract 995, Nijmegen#824750021 (AY266242). S. quitoense Lam.1—Bohs 2873 (UT),Costa Rica (AY266228). S. quitoense Lam.4—Heiser s.n. BohsDNA extract 996, Ecuador: Quitomarket (A Y266243). S. repandum G. Forst.3—Heiser 8215 (IND), Fiji(A Y266229). S. repandum G. Forst.4—Ashley 8627 (IND),Solomon Islands: Malaita (AY266234). S. sessiliorum Dunal3— Dickson 458 (BH) from Whalen 859 (HUT),Peru (AY266261). S. sessiliorum Dunal var. sessiliorum 4—Heiser 8255 (IND), Ecuador: Yanzatza (AY266260). S. stramonifolium Jacq. var. inerme (Dunal) Whalen 4—Pickersgill 154 (IND),Peru: Iquitos (AY266244). S. stramonifolium Jacq. var. inerme (Dunal) Whalen 3—Whalen &Salick 860 (BH),Peru: Pasco, Iscozacin (AY266263). S. vestissimum Dunal3—Dickson 456 (BH) from Plowman 13431 (F),V enezuela (AY266264). S. vestissimum Dun- al4—Movilla s.n. (IND) from Heiser S432a,Colombia: Santa Marta (AY266247). Outgroups: S. abutiloides (Griseb.)Bitter & Lillo 2—RGO S-73 (WTU),BIRM S.0655 (AY266236). S. acerifolium Dunal1—Bohs 2714 (UT),Costa Rica (AY266249). S. capsicoides All.1—Bohs 2451 (UT),Peru (AY266251). S. dulcamara L.2—no voucher, USA(A Y266231). S. jamaicense Mill.2—RGO S-85 (WTU),BIRM S.1209 (AY266239). S. luteoalbum Pers.1—Bohs 2337 (UT), BIRMS.0042 (AY266257). S. mammosum L.2—RGO S-89 (WTU),BIRM S.0983 (AY266232). S. melongena L.2—RGO S-91 (WTU),BIRM S.0657 (AY266240). S. palinacanthum Dunal1—Bohs 3151 (UT),Bolivia (AY266233). S. pseudocapsicum L.2— no voucher, BIRMS.0870 (AY266241). S. robustum Wendl.4—Bohs 3084 (UT),Argentina: Corrientes,Pericho ´n(AY266259). S. sisymbriifolium Lam.1—Bohs 2533 (UT),Argentina (A Y266235). S. stagnale Moric.4—Carvalho 3213 (IND),Brazil: Bahia, Val- enc¸a(AY266262). S. tenuispinum Rusby1—Bohs 2475 (UT),Bolivia (AY266245). S. torvum Sw.1—RGO S-101 (WTU), BIRM S.0839 (AY266246). S. wendlandii Hook. f.1—no voucher, BIRMS.0488 (AY266248). site data(H eiser 1972,1985b, 1987, 1989; Whalen et al. MATERIALSAND METHODS 1981;Whalen and Caruso1983; Bernarde llo et al.1994; Allspecies placed insection Lasiocarpa byWhalen et al. (1981) Bruneau et al.1995). Many ofthese studies were aimed were sampled, including S. stagnale and S. robustum. Solanuma th- atexamining the evolutionary history ofthe Asian dis- eniae isknown only from the type (Symon1985) and no material juncts and the origin and evolution of S.quitoense . De- was available forsampling. Inmost cases, twoaccessions were spite the accumulationof an impressiveamountof sampled fromeach species ofsection Lasiocarpa.Outgroup taxa data,a consensus has not been reached regarding the included ten species from