Edited by K. Kubitzki Volume XV Flowering Plants Eudicots Apiales, Gentianales (except Rubiaceae) Joachim W. Kadereit · Volker Bittrich (Eds.) THE FAMILIES AND GENERA OF VASCULAR PLANTS EditedbyK.Kubitzki Forfurthervolumesseelistattheendofthebookand: http://www.springer.com/series/1306 The Families and Genera of Vascular Plants Edited by K. Kubitzki XV (cid:1) Flowering Plants Eudicots Apiales, Gentianales (except Rubiaceae) Volume Editors: Joachim W. Kadereit (cid:129) Volker Bittrich With 85 Figures Editors JoachimW.Kadereit VolkerBittrich JohannesGutenberg Campinas Universita¨tMainz Brazil Mainz Germany SeriesEditor Prof.Dr.KlausKubitzki Universita¨tHamburg BiozentrumKlein-FlottbekundBotanischerGarten 22609Hamburg Germany TheFamiliesandGeneraofVascularPlants ISBN978-3-319-93604-8 ISBN978-3-319-93605-5 (eBook) https://doi.org/10.1007/978-3-319-93605-5 LibraryofCongressControlNumber:2018961008 #SpringerInternationalPublishingAG,partofSpringerNature2018 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthematerial is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafterdeveloped. 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ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Preface It is with great pleasure that I present a volume with family treatments of the orders ApialesandGentianales(exceptRubiaceae).Animmenseamountofevidencerecently accruedhadtobetakenintoaccounttopresentup-to-datetreatmentsofthesefamilies, including evidence from taxonomic revisions of large groups, new comparative mor- phological studies, but also data from karyological and phytochemical findings and, aboveall,resultsfromanincreasingnumberofmolecularsystematicstudiesthathave changedourconceptsofrelationshipsconsiderably.InthetwolargefamiliesApiaceae andApocynaceae,whichcomprise466and378generarespectively,theestablishment ofaphylogeneticframeworkobviouslyhasbeenacomprehensiveeffortformanyyears by several authors. Although much smaller, the same is true for Loganiaceae where establishment of family limits proved to be particularly difficult. The authors of the Apiaceaetreatmentfounditdifficulttodeviseasingleusablekeytothegeneraofthis family.Therefore,separatekeyswereconstructedforthenativeandnaturalizedgenera in seven major geographic regions of the world. The treatments of Apiaceae and Apocynaceaealsoincludemanyreferencestovariouskeysandtaxonomictreatments of parts of the families that are difficult to find, which makes these treatments even moreuseful. Altogether, the volume contains a wealth of interesting information, and I am greatlyindebtedtoallauthorsfortheirardentdesiretocompletetheircontributions, andtothevolumeeditorswhotookcarethattheprojectwasbroughttoanend.Iam also grateful to all copyright holders who so kindly gave permission to reproduce illustrations published under their responsibility, and to all artists who contributed original illustrations. The artist Bobbi Angell, New York, again deserves our special thanks for the generosity with which she authorized the use of fine illustrations published under her authorship. Doris Franke, Mainz, is gratefully acknowledged for preparingallfiguresforprint. Finally,Ihavegreatpleasureinthankingthecopyeditorofthepresentvolume,Dr. MoniqueDelafontaine,forherdedicatededitorialwork.MywarmthanksalsogotoDr. SabinevonMering,whokindlytookcareofpreparingtheindexofscientificnames.As always, it is a pleasure to acknowledge the cordial collaboration with Dr. Andrea SchlitzbergerfromSpringerVerlag. Hamburg KlausKubitzki 2May2018 v Contents IntroductiontotheOrdersofthis V.BITTRICHandJ.W.KADEREIT ........ 1 Volume Apiaceae G.M.PLUNKETT,M.G.PIMENOV, J.-P.REDURON,E.V.KLJUYKOV, B.-E.VANWYK,T.A.OSTROUMOVA, M.J.HENWOOD,P.M.TILNEY,K.SPALIK, M.F.WATSON,B.-Y.LEE,F.-D.PU, C.J.WEBB,J.M.HART,A.D.MITCHELL, andB.MUCKENSTURM ................... 9 Apocynaceae M.E.ENDRESS, U. MEVE, D.J.MIDDLETON, and S. LIEDE-SCHUMANN ............. 207 Araliaceae G.M.PLUNKETT,J.WEN,P.P.LOWRY II, A.D.MITCHELL,M.J.HENWOOD, andP.FIASCHI ....................... 413 Gelsemiaceae L.STRUWE ............................ 447 Gentianaceae L.STRUWE andJ.S.PRINGLE .......... 453 Griseliniaceae M.O.DILLON ......................... 505 Loganiaceae L.STRUWE,K.L.GIBBONS,B.J.CONN, andT.J.MOTLEY ..................... 511 Myodocarpaceae P.P.LOWRYIIandG.M.PLUNKETT ... 527 Pennantiaceae M.J.POTGIETER ....................... 533 Pittosporaceae R.C.CAROLIN andV.BITTRICH ....... 539 Torricelliaceae G.M.PLUNKETT, Q.-Y. XIANG, P.P. LOWRY II, and G.E. SCHATZ .... 549 Index ........................................ 557 Volumespublishedinthisseries ........................................ 569 vii List of Contributors VolkerBittrich RuaDr.MariodeNucci500,13083-290Campinas,SP,Brazil. [email protected] RogerC.Carolin OakPark,P.O.Box25,Berry2535,NSW,Australia. [email protected] BarryJ.Conn JohnRayHerbarium,SchoolofLifeandEnvironmental Sciences,UniversityofSydney,Sydney,NSW2006,Australia. [email protected] MichaelO.Dillon BotanyDepartment,TheFieldMuseum,1400SouthLakeShore Drive,Chicago,IL60605,USA. mdillon@fieldmuseum.org MaryE.Endress InstituteofSystematicandEvolutionaryBotany,Universityof Zurich,Zollikerstrasse107,8008Zurich,Switzerland. [email protected] PedroFiaschi DepartamentodeBotaˆnica,CCB,UniversidadeFederaldeSanta Catarina,CampusUniversita´rio–Trindade,88040-900 Floriano´polis,SantaCatarina,Brazil. pedrofi[email protected] KerryL.Gibbons RoyalBotanicGardensandDomainTrust,MrsMacquariesRd, Sydney,NSW2000,Australia. [email protected] JenniferM.Hart SchoolofLifeandEnvironmentalSciences,Universityof Sydney,Camperdown,NSW2006,Australia. [email protected] MurrayJ.Henwood SchoolofLifeandEnvironmentalSciences,Universityof Sydney,Camperdown,NSW2006,Australia. [email protected] JoachimW.Kadereit Institutfu¨rOrganismischeundMolekulareEvolutionsbiologie, JohannesGutenberg-Universita¨tMainz,55099Mainz,Germany. [email protected] EugeneV.Kljuykov BotanicalGarden,LomonosovMoscowStateUniversity,1-12 LeninskieGory,119991Moscow,Russia. [email protected] Byoung-YoonLee PlantResourcesDivision,NationalInstituteofBiological Resources,Incheon404-708,RepublicofKorea. [email protected] ix x ListofContributors SigridLiede-Schumann UniversityofBayreuth,Universita¨tstraße30,95440Bayreuth, Germany. [email protected] PorterP.LowryII MissouriBotanicalGarden,P.O.Box299,St.Louis,MO 63166-0299,USA InstitutdeSyste´matique,E´volutionetBiodiversite´(ISYEB), Unite´MixtedeRecherche7205,Muse´umnationald’Histoire naturelle,SorbonneUniversite´s,CP39,57rueCuvier,75231 ParisCEDEX05,France. [email protected] UlrichMeve DepartmentofPlantSystematics,UniversityofBayreuth, Universita¨tstraße30,95440Bayreuth,Germany. [email protected] DavidJ.Middleton SingaporeBotanicGardens,NationalParksBoard,1Cluny Road,Singapore259569,Singapore. [email protected] AnthonyD.Mitchell UniversityofOtago,2RiccartonAvenue,P.O.Box4345, Christchurch8140,NewZealand. [email protected] TimothyJ.Motley DepartmentofBiologicalSciences,OldDominionUniversity, 45thStreet,Norfolk,VA23529,USA BernardMuckensturm 14rueSainte-Anne,98200Mulhouse,France TatianaA.Ostroumova BotanicalGarden,LomonosovMoscowStateUniversity,1-12 LeninskieGory,119991Moscow,Russia. [email protected] MichaelG.Pimenov BotanicalGarden,LomonosovMoscowStateUniversity,1-12 LeninskieGory,119991Moscow,Russia. [email protected] GregoryM.Plunkett CullmanProgramforMolecularSystematics,NewYork BotanicalGarden,2900SouthernBlvd.,Bronx,NY10458-5126, USA. [email protected] MartinJ.Potgieter DepartmentofBiodiversity,UniversityofLimpopo,PrivateBag X1106,Sovenga0727,SouthAfrica. [email protected] JamesS.Pringle RoyalBotanicalGardens,680PlainsRoadWest,Burlington,ON L7T4H4,Canada. [email protected] Fa-DingPu ChengduInstituteofBiology,ChineseAcademyofSciences, P.O.Box416,Chengdu,Sichuan610041,People’sRepublicof China Jean-PierreReduron ViaApia10ruedel’Arsenal,68100Mulhouse,France. [email protected] GeorgeE.Schatz MissouriBotanicalGarden,P.O.Box299,St.Louis,MO 63166-0299,USA. [email protected] ListofContributors xi KrzysztofSpalik DepartmentofMolecularPhylogenetics&Evolution,Facultyof Biology,UniversityofWarsawBiological&ChemicalResearch Centre,101Z˙wirkiiWigurySt.,WarszawaPL-02-096,Poland. [email protected] LenaStruwe DepartmentofEcology,Evolution,&NaturalResources, RutgersUniversity,237ForanHall,59DudleyRoad,New Brunswick,NJ08901,USA. [email protected] PatriciaM.Tilney DepartmentofBotanyandPlantBiotechnology,Universityof Johannesburg,P.O.Box524,AucklandPark2006Johannesburg, SouthAfrica. [email protected] Ben-ErikvanWyk DepartmentofBotanyandPlantBiotechnology,Universityof Johannesburg,P.O.Box524,AucklandPark2006Johannesburg, SouthAfrica. [email protected] MarkF.Watson RoyalBotanicGardenEdinburgh,20aInverleithRow, EdinburghEH35LR,UK. [email protected] ColinJ.Webb TertiaryEducationCommission,Wellington,NewZealand JunWen DepartmentofBotany,NationalMuseumofNaturalHistory, MRC166,SmithsonianInstitution,Washington,DC20013- 7012,USA. [email protected] Qiu-Yun(Jenny)Xiang DepartmentofPlantandMicrobialBiology,NorthCarolina StateUniversity,Raleigh,NC27695-7612,USA. [email protected] Introduction to the Orders of this Volume V. BITTRICH AND J.W. KADEREIT Thepresentvolumeofthisbookseriescompletes – Leaves simple orcompound, petiole usually sheath- the treatment of the Asterids. Asterids are now ing at base; inflorescences or their ultimate units contained in Vols. VI (Cornales, Ericales, 2004), often umbellate; petals usually shorter; stamen fila- VII (Lamiales, 2004), VIII (Asterales, 2007), XIV ments inflexedin bud; stylodia 2–many, ovary infe- (Aquifoliales, Boraginales, Bruniales, Dipsacales, rior; nectary on ovary roof; fruit a schizocarp or Escalloniales, Garryales, Paracryphiales, Sola- drupaceous 5 nales, Icacinaceae, Metteniusaceae, Vahliaceae, 5. Plantsherbaceous,rarelywoody;leavesmostlycom- 2016) and the present volume, which contains pound, rarely entire, usually membraneous; petals the orders Apiales and Gentianales (except usuallyclawedandwithaninflexedtip;ovules2per Rubiaceae). The only families of Asterids not locule,oneabortive;fruitusuallydryandschizocar- treated in the series are Acanthaceae (Lamiales), pic Apiaceae Convolvulaceae (Solanales) and Rubiaceae (Gen- – Plantsmostlyshrubsorsmalltrees;leavessimpleto tianales). compound, usually subcoriaceous to coriaceous; petals usually broadly inserted, rarely clawed, and withoutinflexedtip;ovule1perlocule;fruitusually drupaceous,rarelydryschizocarpic 6 CONSPECTUSOFTHEFAMILIESOFAPIALES 6. Calyx with evident lobes; sepals and petals 5, petal apexwithoutpointedadaxialthickening,stamens5, 1. Plants without secretory canals; leaves simple; nec- carpels2;fruitwithsecretoryvesicleslocatedimme- taryabsentorpresent;fruitfleshy,1-seeded 2 diatelyadjacenttotheendocarp Myodocarpaceae – Plantswithsecretorycanalsthroughout;leavessim- – Calyxwithminutetoobscurelobes,oftenforminga ple or composed; gynoecial nectary mostly present; truncaterim;sepalsandpetals(3–)5(–12),petalapex fruit fleshy or dry, sometimes schizocarpic, 1- to with pointed adaxial thickening, stamens 5 (3– many-seeded 4 numerous), carpels 2–5(–100+); fruit without secre- 2. Petiolenotsheathing;petalsvalvate;nectaryabsent; toryvesicles Araliaceae ovarysuperior,unilocular Pennantiaceae – Petiole (cid:1) sheathing; petals (sub-)imbricate, rarely induplicate-valvate;nectarypresentorabsent;ovary Seven families (about 522 genera, 6000 spp.) are inferior,3-locularatleastinupperpart 3 currentlyincluded:Apiaceae(466/3820),Aralia- 3. Leaves distichous or subopposite, blade minutely ceae(40/1900),Griseliniaceae(1/7),Myodocar- pellucid-punctate; ovary 3-locular above, 1-locular paceae (2 / 17), Pennantiaceae (1 / 4), below Griseliniaceae Pittosporaceae (9 / 250) and Torricelliaceae (s.l. – Leaves spiral, blade not punctate; ovary 3-locular ¼ incl. Aralidiaceae and Melanophyllaceae; 3 / throughout Torricelliaceae 10). The phylogeny currently accepted is (Pen- 4. Leaves simple, petiole not sheathing at base; nantiaceae + (Torricelliaceae + (Griseliniaceae + inflorescences not umbellate; petals 4–40 mm long; (Pittosporaceae + (Araliaceae + (Myodocarpa- stamenfilamentsstraightinbud;stylesimple,ovary ceae+Apiaceae)))))). superior; nectary at base of ovary (rarely absent); Few of the families now included in the fruitacapsuleorberry,rarelywoodyindehiscent Apiales were placed in the order before data Pittosporaceae from DNA sequencing became available. Undis- puted were only Apiaceae + Araliaceae (see #SpringerInternationalPublishingAG,partofSpringerNature2018 1 J.W.Kadereit,V.Bittrich(eds.),FloweringPlants.Eudicots,TheFamiliesandGeneraofVascularPlants15, https://doi.org/10.1007/978-3-319-93605-5_1