Abstract
The cosmopolitan genus Euphorbia is one of the largest plant genera with an extreme phenotypic plasticity ranging from globose succulents to large shrubs and trees adapted to very divergent habitats like forests or deserts. We compiled the current knowledge about the evolution of Euphorbia by reconstructing a molecular phylogeny based on nearly all available internal transcribed spacer (nrITS) sequence data from previous investigations and sequences of newly sampled species. We compared the results with phylogenetic analyses based on a combined data set of nrITS and trnL–trnF intergenic spacer sequences of a subset of taxa. Apart from a few exceptions, the results correspond well to recently published studies based on different molecular markers. The genus Euphorbia is divided into four major clades, of which the clade comprising subgen. Rhizanthium is restricted to Africa and Madagascar, whereas the remaining three clades are distributed over several continents. Our results support the hypothesis that Euphorbia evolved in Africa from progenitors of subgen. Esula.
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Acknowledgments
We thank the curators and staff of the following herbaria kindly sending their material on loan: BISH, BR, F, GOET, HEID, K, MO, NU, P, RSA, TEX, UC and WAG. The following herbaria permitted access to their collections, which is thankfully acknowledged here: B, BOL, BRI, GRA and PRE. The Botanical Gardens of Berlin and Heidelberg, Germany, as well as the Sukkulenten-Sammlung in Zürich, Switzerland, kindly supplied living material. We thank H. Dietrich, W. Ewest, J. Müller, E. Specks (Exotica Nursery), Germany, P. I. Forster, G. & N. Sankowsky, Australia, G. Marx, D. Hardy, South Africa, and M. Machado, Brazil, for the contribution of living plants. P. I. Forster (BRI) generously supported the field trip and herbarium studies of the first author to Queensland. We thank the nature conservation authorities of South Africa, which issued permits for the collection and export of plant material. We thank the two anonymous reviewers for thoughtful comments on the manuscript. The study was financially supported by a grant from the DFG (Deutsche Forschungsgemeinschaft) in 1999.
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N. F. A. Zimmermann and C. M. Ritz contributed equally to the paper.
N. F. A. Zimmermann: Defunct 3 May 2006.
Appendix: Genbank accession number of taxa
Appendix: Genbank accession number of taxa
For species whose markers were sequenced during this study, complete voucher information is given; for species whose sequences were taken from Genbank, accession number and citation are listed. BGJ = Botanical Garden Jena
nrITS sequences
Anthostemamadagascariense Baill., Steinmann & Porter (2002), AF537582;A.sp., Steinmann & Porter (2002), AF537583.
Calycopepluscasuarinoides L.S.Sm., Australia, Queensland, Weipa, Sandy Creek, 10m, 09.07.1993, Forster 13501 & Sankowsky (BRI) AJ534829 AJ534830.
Dichostemmaglaucescens Pierre, Steinmann & Porter (2002), AF537584.
Euphorbiaaaron-rossii A.H. Holmgren & N.H. Holmgren, Steinmann & Porter (2002), AF537396; E.abdelkuri Balf.f., Steinmann & Porter (2002), AF537458; E.acalyphoides Hochst. ex Boiss., Steinmann & Porter (2002), AF537576; E. acanthothamnos Heldr. & Sart. ex Boiss., Steinmann & Porter (2002), AF537554;E.acuta Engelm., Steinmann & Porter (2002), AF537450; E. adiantoides Lam., Steinmann & Porter (2002), AF537395; E. aeruginosa Schweickerdt, Bruyns et al. (2006), AM040756; E. agowensis Hochst. ex Boiss., Steinmann & Porter (2002), AF537419; E. alta Norton, Steinmann & Porter (2002), AF537553;E.amygdaloides L., Steinmann & Porter (2002), AF537544; E. angusta Engelm., Steinmann & Porter (2002), AF537449; E. ankarensis Boiteau, Steinmann & Porter (2002), AF537462; E. ankazobensis Rauh & Hofstätter, Haevermans et al. (2004), AJ508966; E. annamarieae Rauh, Haevermans et al. (2004), AJ508965; E.antiquorum L., Bruyns et al. (2006), AM040757; E. antisyphilitica Zucc., ex cult., Botanical Garden Heidelberg, Germany, Rauh 49959, (BGJ) AJ534791, AJ534792; E. antso Denis, Steinmann & Porter (2002), AF537579; E. aphylla Brouss. ex Wild., Steinmann & Porter (2002), AF537540 (specimen 1); Molero et al. (2002), AF334250, AF334265 (specimen 2); Canary Islands, Tenerife, Punta de Teno, April 1994, Zimmermann 11/94 (BGJ) AJ534827, AJ534828 (specimen 3); E. appariciana Rizzini, Steinmann & Porter (2002), AF537455 (specimen 1); Brazil, Bahia, E of Morro do Chapéu, 12.03.1989, Supthut 8908 (ZSS) AJ534783, AJ534784 (specimen 2); E. arbuscula Balf. f., Steinmann & Porter (2002), AF537496; E. arceuthobioides Boiss., Bruyns et al. (2006), AM040758; E. articulata Burm., Steinmann & Porter (2002), AF537446; E. atrispina N.E.Br., Steinmann & Porter (2002), AF537568; E. atropurpurea Brouss. ex Willd., Steinmann & Porter (2002), AF537542; E. attastoma Rizzini, Steinmann & Porter (2002), AF537511; E. atoto G. Forst. Australia, Queensland, Wongaling Beach, 01.12.1999, Zimmermann 49/99 (BGJ) AJ534807, AJ534808; E. balsamifera Aiton ssp.adensis (Deflers) Bally, Steinmann & Porter (2002), AF537571; E. beharensis Leandri, Haevermans et al. (2004), AJ508983; E. bemarahaensis Rauh & Mangelsdorff, Haevermans unpubl., AJ508984; E. bicolor Engelm. & A. Gray, Steinmann & Porter (2002), AF537386; E. bifurcata Engelm., Steinmann & Porter (2002), AF537434; E. bilobata Engelm., Steinmann & Porter (2002), AF537435; E. biselegans Bruyns, Steinmann & Porter (2002), AF537470 (specimen 1); Tanzania, Iringa, N of Malolo, Specks 699 (BGJ) AJ534759, AJ534760 (specimen 2); E. boissierii Baill., Haevermans et al. (2004), AJ508963; E. boophthona C.A. Gardner, Steinmann & Porter (2002), AF537515 (specimen 1); Australia, Queensland, E of Quilpie, 22.11.1999, Zimmermann 25/99 (BGJ) AJ534811, AJ534812 (specimen 2); E. bovinii Boiss., Haevermans et al. (2004), AJ508962; E.brachyphylla Denis, Haevermans et al. (2004), AJ508961; E.bracteata Jacq., Steinmann & Porter (2002), AF537489; E. brunellii Chiov., Steinmann & Porter (2002), AF537486; E. bruynsii L.C. Leach, Bruyns et al. (2006); E. bulbispina Rauh, Haevermans et al. (2004), AJ508960; E. bupleurifolia Jacq., South Africa, Eastern Cape, Grahamstown, 06.03.1998, Zimmermann 12/98 (BGJ) AJ536017, AJ536018; E. burmannii E. Mey ex. Boiss., Bruyns et al. (2006), AM040760; E. calcarata (Schltdl.) V. W. Steinm., Steinmann & Porter (2002), AF537492; E. calcicola Fern., Steinmann & Porter (2002), AF537385; E. californica Benth., Steinmann & Porter (2002), AF537377; E. calyculata Kunth, Steinmann & Porter (2002), AF537524; E. calyptrata Coss. & Kralik, Steinmann & Porter (2002), AF537549; E. capmanambatoensis Rauh, Steinmann & Porter (2002), AF537476 (specimen 1); Haevermans et al. (2004), AJ508959 (specimen 2); E. caput-medusae L., South Africa, Western Cape, S of Camps Bay, 05.03.2000, Zimmermann 30/00 (BGJ), AJ535533, AJ535555; E. carunculata Waterf. Steinmann & Porter (2002), AF537447; E. cassythoides Boiss., Steinmann & Porter (2002), AF537387; E. cedrorum Rauh & Hebding, Haevermans et al. (2004), AJ508958 (specimen 1); ex cult., Botanical Garden Heidelberg, no. 140124, Germany, Rauh (J.B.H.)59367a (HEID) AJ534757, AJ534758 (specimen 2); E.celastroides Boiss., Hawaiian Islands, Kauai, Kilauea Point, March 1999, Dietrich s.n. (BGJ), AJ534799, AJ534800; E. ceroderma I.M. Johnst., Steinmann & Porter (2002), AF537389; E. cestrifolia Kunth, Steinmann & Porter (2002), AF537521; E. cheirolepis Fisch & C.A. Mey, Steinmann & Porter (2002), AF537424; E.chersonesa Huft, Steinmann & Porter (2002), AF537436; E.clava Jacq., Bruyns et al. (2006), AF537569; E.colletioides Benth., Steinmann & Porter (2002), AF537405; E. colligata V.W. Steinm., Steinmann & Porter (2002), AF537493; E. comosa Vell., Steinmann & Porter (2002), AF537503; E. contorta L.C. Leach, Bruyns et al. (2006), AM040761; E. corniculata R.A. Dyer, Bruyns et al. (2006), AM040762; E. cremersii Rauh & Razaf., Haevermans et al. (2004), AJ508957; E. croizatii (Hurus) Kitag., Haevermans et al. (2004), AJ508956; E. crossadenia Pax & K. Hoffm., Steinmann & Porter (2002), AF537451; E. crotonoides Boiss., Steinmann & Porter (2002), AF537578; E. cubensis Boiss. Steimann et al. (2007), EF653254 (specimen 1); Cuba, Pinar del Rio, Cuabales, San Juan de Saqua, 22.02.2001, Hellwig 22/2/28 (BGJ), AJ534775, AJ534776 (specimen 2); E.cupularis Boiss., South Africa, Transvaal, Njele Poort, 1978, Hardy 5115 (PRE), AJ534765, AJ534766; E.cyathophora Murray, Cyprus, Limassol, Kolossi Castle, 30.10.1995, Kuschel 644 (BGJ), AJ534797, AJ534798; E. cylindrifolia Marn.-Lapost. & Rauh, Haevermans et al. (2004), AJ508955; E. cymbifera (Schltdl.) Steinm., Steinmann & Porter (2002), AF537491; E. finkii (Boiss.) V.W. Steinm., Steinmann & Porter (2002), AF537520; E. cyparissias L., Germany, Thuringia, S of Uhlstädt, 13.04.1998, Zimmermann 74/98 (BGJ), AJ534823, AJ534824; E. decaryi Guillaumin, Madagascar, Vinanibé near Fort Dauphin, Rauh 68100 (HEID), AJ534751, AJ534752; E. decidua P.R.O. Bally & L.C. Leach, Bruyns et al. (2006), AM040763; E. decorsei Drake, Haevermans unpubl., AJ508985; E. degeneri Sherff., Steinmann & Porter (2002), AF537444; E.delicatula Boiss., Steinmann & Porter (2002), AF537393; E. dendroides L., Steinmann & Porter (2002), AF537539; E. denisii Oudejans, Steinmann & Porter (2002), AF537497; E. dentata Michx., ex cult., Botanical Garden Jena, Germany, Zimmermann 102/00 (BGJ), AJ534795, AJ534796; E. depauperata Hochst. ex A. Rich, Steinmann & Porter (2002), AF537556; E. didiereoides Denis ex Leandri, Haevermans et al. (2004), AJ508954 (specimen 1); ex cult., Botanical Garden Heidelberg No. 140087, Germany, Rauh 71949 (HEID), AJ534741, AJ534742; E. discolor Ledeb., Steinmann & Porter (2002), AF537547; E. dregeana E. Mey. ex Boiss., South Africa, Northern Cape, S of Vioolsdrif, 01.03.2000, Zimmermann 20/00 (BGJ), AJ534724, AJ534734; E.drupifera Thonn., Steinmann & Porter (2002), AF537480; ex cult., Botanical Garden Heidelberg, Germany, Rauh 9610 (BGJ), AJ534727, AJ534737; E. eanophylla Croizat, Steinmann & Porter (2002), AF537498; E.echinulata (Stapf) Bruyns, Tanzania, Iringa, between Madibira and Igawa, Specks 718 (BGJ),AJ534767, AJ534768; E. eduardoi L.C. Leach, Bruyns et al. (2006), AM040765; E. eglandulosa V.W. Steinm., Steinmann & Porter (2002), AF537394; E. elata Brandegee, Steinmann & Porter (2002), AF537495; E. elliotii Leandri, Steinmann & Porter (2002), AF537478 (specimen 1); Bruyns et al. (2006), AM040766 (specimen 2); E.epicyparissias (E. Mey. ex Klotzsch & Garcke) Boiss., South Africa, Kwazulu/Natal, E of Sani Pass, 18.03.2000, Zimmermann 70/00 (BGJ), AJ534825, AJ534826; E. epiphylloides Kurz, Steinmann & Porter (2002), AF537484; E. equisetiformis A. Stewart, Steinmann & Porter (2002), AF537388; E. eremophila A. Cunn., Steinmann & Porter (2002), AF537423; E. eriantha Benth., Steinmann & Porter (2002), AF537440; E.esculenta Marloth, Steinmann & Porter (2002), AF537575; E. espinosa Pax, Steinmann & Porter (2002), AF537416;E. esula L., Steinmann & Porter (2002), AF537546; E. exilis L.C. Leach, Bruyns et al. (2006), AM040767; E. exstipulata Engelm., Steinmann & Porter (2002), AF537433; E. fianarantsoae Ursch & Leandri, Ritz et al. unpubl., AJ535538, AJ535560; E. filiflora Marloth, Bruyns et al. (2006), AM040768; E. fulgens Karw. ex Klotzsch, Steinmann & Porter (2002), AF537404; E. gariepina Boiss., Bruyns et al. (2006), AM040769; E. genistoides P.J. Bergius, Bruyns et al. (2006), AM040770; E. gentryi V.W. Steinm. & T.F. Daniel, Steinmann & Porter (2002), AF537406; E. germainii Phil., Steinmann & Porter (2002), AF537499; E. geroldii Rauh, Steinmann & Porter (2002), AF537475; E. glanduligera Pax, Steinmann & Porter (2002), AF537426; E.globosa (Haw.) Sims, South Africa, Eastern Cape, S of Addo, 07.03.1998, Zimmermann 8/98 (BGJ), AJ535535, AJ535557; E. goetzei Pax, Steinmann & Porter (2002), AF537413; E. gollmeriana Klotzsch ex Boiss., Steinmann & Porter (2002), AF537501; E. gottlebei Rauh, Steinmann & Porter (2002), AF537459; E.gradyi V.W. Steinm. & Ram.-Roa, Steinmann & Porter (2002), AF537407; E. graminea Jacq., Steinmann & Porter (2002), AF537410; E. grandicornis Goebel, Bruyns et al. (2006), AM040771; E. graniticola L.C. Leach, Bruyns et al. (2006), AM040772; E. gregaria Marloth, Steinmann & Porter (2002), AF537527; E. griseola Pax, Bruyns et al. (2006), AM040773 (specimen 1); Ritz et al. unpubl., AJ535536, AJ535558 (specimen 2); E.guatemalensis Standl. & Steyerm., Steinmann & Porter (2002), AF537408; E. guerichiana Pax ex Engl., Steinmann & Porter (2002), AF537415 (specimen 1); South Africa, Northern Province, Zoutpan, W of Waterpoort, 22.03.2000, Zimmermann 82/00 (BGJ), AJ534813, AJ534814 (specimen 2); E.gymnoclada Boiss., Steinmann & Porter (2002), AF537456; E. gymnonota Urb., Steinmann & Porter (2002), AF537507; E.haeleeleana Herbst, Steinmann & Porter (2002), AF537514; E. hallii R.A. Dyer, Steinmann & Porter (2002), AF537573; E. hamata Sweet, Bruyns et al. (2006), AM040774; E. hedyotoides N.E.Br., Steinmann & Porter (2002), AF537460 (specimen 1); Madagascar, Ambovombe, Rauh 72073 (HEID), AJ534755, AJ534756 (specimen 2); E. helenae Urb. Steimann et al. (2007), EF653255; E. helioscopia L., Bruyns et al. (2006), AM040775; E. heterodoxa Müll.Arg., Steinmann & Porter (2002), AF537500 (specimen 1); ex cult., Botanical Garden Berlin-Dahlem no. 078-14-96-40, Germany, (BGJ), AJ534777, AJ534778 (specimen 2); E.heterophylla L., Steinmann & Porter (2002), AF537429; E. hoffmanniana Boiss., Steinmann & Porter (2002), AF537508; E. hormorrhiza Radcl.-Sm., Steinmann & Porter (2002), AF537431; E. horombensis Ursch & Leandri, Haevermans et al. (2004), AJ508971; E. hypericifolia L., Steinmann & Porter (2002), AF537443; E.hypogaea Marloth, Ritz et al. (2003) AJ535534, AJ535556; E. horrida Boiss., South Africa, Eastern Cape, Kleinpoort, 13.08.1997, Zimmermann 317/97 (BGJ), AJ535525, AJ535547; E. iharanae Rauh, Steinmann & Porter (2002), AF537477; E. ingens E. Mey. ex Boiss., South Africa, Mpumalanga, E of Burgersfort, 31.07.1997, Zimmermann 274/97 (BGJ), AJ534726, AJ534736; E. innocua L.C. Wheeler, Steinmann & Porter (2002), AF537380; E. insulana Vell., Steinmann & Porter (2002), AF537411; E. intisy Drake, Haevermans unpubl., AJ508986; E. ipecacuanhae L., Steinmann & Porter (2002), AF537397; E. jaliscensis B.L. Rob. & Greenm., Steinmann & Porter (2002), AF537442; E. jansenvillensis Nel., Ritz et al. (2003) AJ535524, AJ535546; E. juglans Compton, Ritz et al. (2003), AJ535529, AJ535551 (specimen 1), AJ535530, AJ535552 (specimen 2); E. juttae Dinter, Steinmann & Porter (2002), AF537418; E. kamponii Rauh & H. Petignat, Haevermans et unpubl., AJ508987; E. knuthii (N.E.Br.) L.C. Leach, Bruyns et al. (2006), AM040776; E.kraussiana Bernh. ex Krauss, Steinmann & Porter (2002), AF537548; E. lacera Boiss., Steinmann & Porter (2002), AF537441; E. lactiflua Phil., Steinmann & Porter (2002), AF537528; E. lagunensis Huft, Steinmann & Porter (2002), AF537379; E. lagunillarum Croizat, Steinmann & Porter (2002), AF537502;E. lathyris L., Steinmann & Porter (2002), AF537549; E. laurifolia Lam., Steinmann & Porter (2002), AF537509; E. leistneri R.H. Archer, Bruyns et al. (2006), AM040777; E. leucocephala Lotsy, Steinmann & Porter (2002), AF537381; E. lignosa Marloth, ex cult., Botanical Garden Heidelberg, Germany, Rauh 19624 (BGJ), AJ534723, AJ534733; E. limpopoana L.C. Leach ex S. Carter, Bruyns et al. (2006), AM040778; E. lindenii (S. Carter) Bruyns, Steinmann & Porter (2002), AF537473; E. lomelii V.W. Steinm., Steinmann & Porter (2002), AF537490 (specimen 1); Mexico, Sinaloa, Mazatlán, 10.06.1970, Rauh 50464 (BGJ), AJ534815, AJ534816 (specimen 2); E. longifolia Lam., Steinmann & Porter (2002), AF537558; E. longituberculosa Boiss., Steinmann & Porter (2002), AF537577; E. lophogona Lam., Haevermans et al. (2004), AJ508972 (specimen 1); ex cult., Botanical Garden Heidelberg No. 140013, Germany, Rauh 22116 (HEID), AJ534747, AJ534748 (specimen 2); E.lugardiae (N.E.Br.) Bruyns, Bruyns et al. (2006), AM040799; E. lumbricalis L.C. Leach, Bruyns et al. (2006), AM040779; E. lydenburgensis Schweick. & Letty, Bruyns et al. (2006), AM040755; E. macropus Boiss., Steinmann & Porter (2002), AF537378; E. maculata L., Germany, Thuringia, Jena, 13.07.2000, Zimmermann 101/00 (BGJ), AJ534801, AJ534802; E. macvaughii Carvajal & Lomeli, Steinmann & Porter (2002), AF537382; E. magnifica (E.A. Bruce) Bruyns, Steinmann & Porter (2002), AF537472; E.mahabobokensis Rauh, Steinmann & Porter (2002), AF537522; E. mahafalensis Denis, Haevermans et al. (2004), AJ508973; E. malevola L.C. Leach, Bruyns et al. (2006), AM040780; E. marginata Pursh, ex cult., Botanical Garden Jena, Germany, Zimmermann 103/00 (BGJ), AJ534787, AJ534788; E. mauritanica L., Steinmann & Porter (2002), AJ534821 (specimen 1); South Africa, Northern Cape, N of Pofadder, 29.02.2000, Zimmermann 12/00 (BGJ), AJ534821, AJ534822 (specimen 2); E. medicaginea Boiss., Steinmann & Porter (2002), AF537535; E. meenae S. Carter, Steinmann & Porter (2002), AF537483; E. megalatlantica Baill., Steinmann & Porter (2002), AF537536; E. meloformis Aiton, Ritz et al. (2003) AJ535517; AJ535542; E.meloformis fma. magna, Ritz et al. (2003), AJ535521, AJ535543; E. meuleniana O. Schwartz, Steinmann & Porter (2002), AF537572; E. milii Des Moul., Steinmann & Porter (2002), AF537461 (specimen 1); Haevermans et al. (2004), AJ508974 (specimen 2); E. millotii Ursch & Leandri, Steinmann & Porter (2002), AF537463 (specimen 1); ex cult., Botanical Garden Heidelberg, Germany, Rauh 8608 (HEID), AJ534749, AJ534750; E.misella S. Watson, Steinmann & Porter (2002), AF537384; E. misera Benth., Steinmann & Porter (2002), AF537383; E. mlanjeana L.C. Leach, Bruyns et al. (2006), AM040781; E. monadenioides M.G. Gilbert,Ethiopia, Vlk 344 (BGJ), AJ534729, AJ534739; E. monteiroi Hook.f., Steinmann & Porter (2002), AF537563; E. multifolia A.C. White, R.A. Dyer & B. Sloane, Bruyns et al. (2006), AM040782; E. munizii Borhidi, Steimann et al. (2007), EF653256; E. myrsinites L., Steinmann & Porter (2002), AF537551; E. namuskluftensis L.C. Leach, Steinmann & Porter (2002), AF537562; E. neogossweileri Bruyns, Steinmann & Porter (2002), AF537471 (specimen 1); Angola, Benguella, Lavranos s.n.; ex cult. International Succulent Institute (ISI 1261); leg. Forster 25311 (BRI), AJ534763, AJ534764 (specimen 2); E. neopedunculata Bruyns, Bruyns et al. (2006), AM040800; E. neospinescens Bruyns, Tanzania, Mbeya, Lake Rukwa, Illoma Mt., Specks 1373 (BGJ), AJ534761, AJ534762; E. neriifolia L., Bruyns et al. (2006), AM040783; E. nivulia Buch.-Ham., Steinmann & Porter (2002), AM040784; E. nutans Lag., Canary Islands, Gran Canaria, Maspalomas, Faro II, 24.02.1996, Kuschel 646 (BGJ), AJ534803, AJ534804; E. oaxacana B.L. Rob. & Greenm., Steinmann & Porter (2002), AF537373; E. obesa Hook.f., Ritz et al. (2003), AJ535519, AJ535519; E. oblongata Griseb., Steinmann & Porter (2002), AF537555; E. oncoclada Drake, Steinmann & Porter (2002) (specimen 1); AF537468, Haevermans et al. (2004), AJ508969 (specimen 2); E. ocymoidea L., Steinmann & Porter (2002), AF537392; E.omariana M.G. Gilbert, Steinmann & Porter (2002), AF537560; E. orthoclada Baker,Madagascar, Androy, 10.10.1998, Rauh 70626 (HEID), AJ534819, AJ534820; E. oxystegia Boiss., Bruyns et al. (2006), AM040785; E. pachysantha Baill., Haevermans et al. (2004), AJ508976; E. pachypodioides Boiteau, Madagascar, Ankarana, 25.03.1991, Rauh 72072 (HEID), AJ534753, AJ534754; E. panchganiensis Blatt. & McCann, Steinmann & Porter (2002), AF537375; E. papillosa A.St.-Hil., Steinmann & Porter (2002), AF537510; E.parvicaruncula D.C. Hassall, Australia, Queensland, S of Boulia, 25.11.1999, Zimmermann 30/99 (BGJ), AJ534809, AJ534810; E. parvicyathophora Rauh, Haevermans et al. (2004), AJ508977; E. pauliani Ursch & Leandri, Haevermans et al. (2004), AJ508978; E. pedilanthoides Denis, ex cult., Botanical Garden Heidelberg No. 142066, Germany, Rauh 66824 (HEID) AJ534745, AJ534746; E. pentadactyla Griseb. Steinmann & Porter (2002), AF537428; E. peperomioides Boiss., Steinmann & Porter (2002), AF537523; E. peplus L., Steinmann & Porter (2002), AF537532; E. perrierii Drake, Steinmann & Porter (2002), AF537464; E. pervilleana Baill., Steinmann & Porter (2002), AF537518; E. petiolata Banks. & Sol., Steinmann & Porter (2002), AF537422; E.phosphorea Mart., Steinmann & Porter (2002), AF537512; E. phylloclada Boiss., Steinmann & Porter (2002), AF537427; E. pilosa L., Steinmann & Porter (2002), AF537557; E. pirottae Terrac., Steinmann & Porter (2002), AF537417; E. piscatoria Aiton, ex cult., Botanical Garden Berlin-Dahlem, No. 075-01-96-10, AJ534817, AJ534818; E. plagiantha Drake, Haevermans unpubl., AJ508988; E. platyclada Rauh, Steinmann & Porter (2002), AF537421; E. platycephala Pax, Steinmann & Porter (2002), AF537561; E. plumerioides Teijsm. ex Hassk., Steinmann & Porter (2002), AF537513 (specimen 1); Australia, Queensland, Kennedy Hill Gorge, 21.06.1989, Forster 5402 (BRI), AJ534769, AJ534770 (specimen 2); E. podocarpifolia Urb., Steimann et al. (2007), EF653257; E. poeppigii (Klotzsch & Garcke) Boiss., Bolivia, La Paz, Nor Yungas, Cerro Uchimachi se Coroico, J. Müller 7040 (BGJ), AJ534793, AJ534794; E. poissonii Pax, Steinmann & Porter (2002), AF537482; E. polygona Haw., Bruyns et al. (2006), AJ535525; E. primulifolia Baker, Steinmann & Porter (2002), AF537466; E. prostrata Aiton, Steinmann & Porter (2002), AF537445, E. pseudoglobosa Marloth, Ritz et al. (2003) AJ535527; AJ535528 (specimen 1); AJ535549, AJ535550 (specimen 2); E. pseudograntii Bruyns, Steinmann & Porter (2002), AF537469; E. pteroneura A. Berger, Steinmann & Porter (2002), AF537504; E.pulcherrima Willd. ex Klotzsch, Steinmann & Porter (2002), AF537432; E. punicea Sw., Steinmann & Porter (2002), AF537516; E. quadrata Nel, Bruyns et al. (2006), AM040786; E. quartziticola Leandri, Haevermans et al. (2004), AJ508979 (specimen 1); Madagascar, Ucal, Itremo, 24.10.1992, Rauh 73705 (HEID), AJ534743, AJ534744 (specimen 2); E. radians Benth., Steinmann & Porter (2002), AF537438; E. ramulosa L.C. Leach, Bruyns et al. (2006), AM040787; E. regis-jubae Webb & Berthel., Steinmann & Porter (2002), AF537541; E. rhombifolia Boiss., Steinmann & Porter (2002), AF537414 (specimen 1); Bruyns et al. (2006), AM040764 (specimen 2); E. robivelonae Rauh, Haevermans et al. (2004), AJ508980; E. robusta (Engelm.) Small, Steinmann & Porter (2002), AF537533; E. rossiana Pax, Steinmann & Porter (2002), AF537374; E.rossii Rauh & Buchloh, Steinmann & Porter (2002), AF537465; E. rubella Pax, Steinmann & Porter (2002), AF537487; E. rzedowskii McVaugh, Steinmann & Porter (2002), AF537399; E. sarcodes Boiss., Steinmann & Porter (2002), AF537454 (specimen 1); Brazil, Bahia, Morro do Chapéu, 25.11.2000, Machado s.n. (BGJ), AJ534785, AJ534786; (specimen 2); E. sarcostemmoides J.H. Willis, Australia, Queensland, W of Quilpie, 22.11.99, Zimmermann 23/99 (BGJ), AJ534771, AJ534772; E. scatorhiza S. Carter, Steinmann & Porter (2002), AF537420; E. schimperi C. Presl, Steinmann & Porter (2002), AF537537; E. schizolepis F. Muell. ex Boiss., Australia, Western Australia, Packsaddle Plain, Kununurra, 04.03.1993, Mitchell 2851 (BRI), AJ534805, AJ534806; E. schlechtendalii Boiss., ex cult., Botanical Garden Heidelberg, Germany, Rauh s.n. (BGJ), AJ534789, AJ534790; E. schoenlandii Pax, Bruyns et al. (2006), AM040788; E. scutiformis V.W. Steinm. & P.E. Berry, Cuba, Sancti Spíritus, Banao, 300-500m, 20.11.1975, Alvarez, Areces, Bisse, Dietrich & Lepper 28763 (BGJ), AJ534773 AJ534774; E. sebsebei M.G. Gilbert, Ethiopia, Sidamo, Yavello, Vlk 326 (BGJ), AJ534730, AJ534740; E. segoviensis Boiss., Steinmann & Porter (2002), AF537400; E. sessilifolia Klotzsch ex Boiss., Steinmann & Porter (2002), AF537452; E. sinaloensis Brandegee, Steinmann & Porter (2002), AF537401; E. silenifolia (Haw.) Sweet, South Africa, Eastern Cape, W of Grahamstown, 05.03.1998, Zimmermann 9/98 (BGJ), AJ534721, AJ534731; E. sipolisii N.E.Br., Steinmann & Porter (2002), AF537517 (specimen 1); Brazil, Minas Gerais, Santa Barbara, Augusto de Lima, 06.08.1988, Eggli 1103 (ZSS), AJ534779, AJ534780 (specimen 2); E. sonorae Rose, Steinmann & Porter (2002), AF537402; E. spathulata Lam., Steinmann & Porter (2002), AF537552; E. sphaerorhiza Benth., Steinmann & Porter (2002), AF537412; E.squarrosa Haw., Bruyns et al. (2006), AM040789; E. stapelioides A.C. White, R.A. Dyer & B. Sloane, Bruyns et al. (2006), AM040790; E. stellata Willd., South Africa, Eastern Cape, Motherwell, N of Port Elizabeth, 12.08.1997, Zimmermann 313/97 (BGJ), AJ534725, AJ534735; E. stenoclada Baill., Haevermans unpubl., AJ508991 (specimen 1); Bruyns et al. (2006), AM040791 (specimen 2); E. stenophylla Boiss., Steinmann & Porter (2002), AF537529; E. stolonifera Marloth, Bruyns et al. (2006), AM040792; E. stricta L., Steinmann & Porter (2002), AF537559; E. strigosa Hook. & Arn., Steinmann & Porter (2002), AF537439; E. subpeltata S. Watson, Steinmann & Porter (2002), AF537376; E. subpeltatophylla Rauh, Haevermans unpubl., AJ508992; E. subsalsa Hiern, Bruyns et al. (2006), AM040793; E. succedanea L.C. Wheeler, Steinmann & Porter (2002), AF537403; E. sudanica A. Chev., ex cult., Botanical Garden Berlin-Dahlem No. 221-04-94-20, AJ534728, AJ534738; E. suffulta Bruyns, Bruyns et al. (2006), AM040794; E. susannae Marloth, Ritz et al. (2003), AJ535531, AJ535532 (specimen 1); AJ535553, AJ535554 (specimen 2); E. symmetrica A.C. White, R.A. Dyer & B. Sloane, Ritz et al. (2003), AJ535518, AJ535539; E. tannensis Spreng., Steinmann & Porter (2002), AF537425; E. tehuacana (Brandegee) V.W. Steinm., Steinmann & Porter (2002), AF537488; E. teke Schweinf. ex Pax, Steinmann & Porter (2002), AF537485; E. tetraptera Baker, Steinmann & Porter (2002), AF537526; E.thinophila Phil., Steinmann & Porter (2002), AF537530 (specimen 1); Chile, N of Huasco, 01.11.00, Ewest 03-11-00 (BGJ), AJ534781, AJ534782 (specimen 2); E. thouarsiana Baill., Steinmann & Porter (2002), AF537474; E. tirucalli L., Steinmann & Porter (2002), AF537479; E. tithymaloides L., Steinmann & Porter (2002), AF537494; E. torrei (L.C. Leach) Bruyns, Bruyns et al. (2006), AM040801; E. trichadenia Pax, South Africa, Gauteng, Soutpan, NW of Pretoria, February 1998, Hardy s.n. (BGJ), AJ534722, AJ534732; E.trichotoma Kunth, Steinmann & Porter (2002), AF537534; E. tuberosa L., Steinmann & Porter (2002), AF537570 (specimen 1); Bruyns et al. (2006), AM040795 (specimen 2); E. tubiglans Marloth ex R.A. Dyer, Ritz et al. (2003), AJ535522, AJ535523 (specimen 1); AJ535544, AJ535545 (specimen 2); E. tulearensis Rauh, Haevermans et al. (2004), AJ508981; E. turbiniformis Chiov., Ritz et al. unpubl., AJ535537, AJ535559; E.turczaninowii Kar. & Kir., Steinmann & Porter (2002), AF537543; E. umbelliformis (Urb. & Ekman) V.W. Steinm. & P.E. Berry, Steimann et al. (2007), EF653253; E. unicornis R.A. Dyer, Bruyns et al. (2006), AM040796; E. usambarica Pax, Steinmann & Porter (2002), AF537538; E. vajravelui Binojk. & N.P. Balakr., Bruyns et al. (2006), AM040797; E. valida N. E. Br., Ritz et al. (2003), AJ535520, E. viguieri Denis, Haevermans et al. (2004), AJ508982; E. waterbergensis R.A. Dyer, Bruyns et al. (2006), AM040798; E.weberbaueri Mansf., Steinmann & Porter (2002), AF537519; E. whitei L.C. Wheeler, Steinmann & Porter (2002), AF537391; E. xylophylloides Brongn. ex Lem., Steinmann & Porter (2002), AF537467; E. zonosperma Müll.Arg., Steinmann & Porter (2002), AF537430.
Neoguillauminiacleopatra (Baill.) Croizat New Caledonia, Mount Dzumac, 700m, 05.12.1980, Morat 6651 (BR), AJ534831, AJ534832
trnL-trnF intergenic spacer sequences
Calycopepluscasuarinoides L.S.Sm. Australia, Queensland, Weipa, Sandy Creek, 10m, 09.07.1993, Forster 13501 & Sankowsky (BRI), FN423771 .
Euphorbiaantisyphilitica Zucc. ex cult., Botanical Garden Heidelberg, Germany, Rauh 49959, (BGJ), FN423716; E. appariciana Rizzini, Brazil, Bahia, E of Morro do Chapéu, 12.03.1989, Supthut 8908 (ZSS), FN423717 (specimen 2); E. aphylla Brouss. ex Willd., Canary Islands, Tenerife, Punta de Teno, April 1994, Zimmermann 11/94 (BGJ), FN423718 (specimen 3); E. atoto G. Forst. Australia, Queensland, Wongaling Beach, 01.12.1999, Zimmermann 49/99 (BGJ), FN423719; E. boophthona C.A. Gardner Australia, Queensland, E of Quilpie, 22.11.1999, Zimmermann 25/99 (BGJ), FN423720 (specimen 2); E. bupleurifolia Jacq., South Africa, Eastern Cape, Grahamstown, 06.03.1998, Zimmermann 12/98 (BGJ), FN423721; E. caput-medusae L. South Africa, Western Cape, S of Camps Bay, 05.03.2000, Zimmermann 30/00 (BGJ), FN423722; E. cedrorum Rauh & Hebding ex cult., Botanical Garden Heidelberg No. 140124, Germany, Rauh (J.B.H.) 59367a (HEID), FN423723 (specimen 2); E. celastroides Boiss., Hawaiian Islands, Kauai, Kilauea Point, March 1999, Dietrich s.n. (BGJ), FN423724; E. cubensis Boiss., Cuba, Pinar del Rio, Cuabales, San Juan de Saqua, 22.02.2001, Hellwig 22/2/28 (BGJ), FN423725 (specimen 2); E. cupularis Boiss., South Africa, Transvaal, Njele Poort, 1978, Hardy 5115 (PRE), FN423726; E. cyathophora Murray, Cyprus, Limassol, Kolossi Castle, 30.10.1995, Kuschel 644 (BGJ), FN423727; E. cyparissias L., Germany, Thuringia, S of Uhlstädt, 13.04.1998, Zimmermann 74/98 (BGJ), FN423728; E. decaryi Guillaumin, Madagascar, Vinanibé near Fort Dauphin, Rauh 68100 (HEID), FN423729; E. dentata Michx., ex cult., Botanical Garden Jena, Germany, Zimmermann 102/00 (BGJ), FN423730; E. didiereoides Denis ex Leandri, ex cult., Botanical Garden Heidelberg No. 140087, Germany, Rauh 71949 (HEID), FN423731 (specimen 2); E. dregeana E. Mey. ex Boiss., South Africa, Northern Cape, S of Vioolsdrif, 01.03.2000, Zimmermann 20/00 (BGJ), FN423732; E. drupifera Thonn., ex cult., Botanical Garden Heidelberg, Germany, Rauh 9610 (BGJ), (specimen 2), FN423733; E. echinulata (Stapf) Bruyns, Tanzania, Iringa, between Madibira and Igawa, Specks 718 (BGJ), FN423734; E. epicyparissias (E. Mey. ex Klotzsch & Garcke) Boiss., South Africa, Kwazulu/Natal, E of Sani Pass, 18.03.2000, Zimmermann 70/00 (BGJ), FN423735; E. fianarantsoae Ursch & Leandri, Ritz et al. unpubl., AJ575321; E. globosa (Haw.) Sims, South Africa, Eastern Cape, S of Addo, 07.03.1998, Zimmermann 8/98 (BGJ), FN423736; E. griseola Pax, Ritz et al. unpubl., AJ575317 (specimen 2); E. guerichiana Pax ex Engl., South Africa, Northern Province, Zoutpan, W of Waterpoort, 22.03.2000, Zimmermann 82/00 (BGJ), FN423737 (specimen 2); E. hedyotoides N.E.Br., Madagascar, Ambovombe, Rauh 72073 (HEID), FN423738 (specimen 2); E. heterodoxa Müll.Arg., ex cult., Botanical Garden Berlin-Dahlem No. 078-14-96-40, Germany, (BGJ), FN423739 (specimen 2); E. horrida Boiss., South Africa, Eastern Cape, Kleinpoort, 13.08.1997, Zimmermann 317/97 (BGJ), FN423740; E. ingens E.Mey. ex Boiss., South Africa, Mpumalanga, E of Burgersfort, 31.07.1997, Zimmermann 274/97 (BGJ), FN423741; E. jansenvillensis Nel, Ritz et al. (2003), AJ575344; E. juglans Compton, Ritz et al. (2003), AJ575346 (specimen 1); AJ57534 (specimen 2); E. lignosa Marloth, ex cult., Botanical Garden Heidelberg, Germany, Rauh 19624 (BGJ), FN423742; E. lomelii V.W. Steinm., Mexico, Sinaloa, Mazatlán, 10.06.1970, Rauh 50464 (BGJ), FN423743 (specimen 2); E. lophogona Lam., ex cult., Botanical Garden Heidelberg No. 140013, Germany, Rauh 22116 (HEID), FN423744, (specimen 2); E. maculata L., Germany, Thuringia, Jena, 13.07.2000, Zimmermann 101/00 (BGJ), FN423745; E. marginata Pursh, ex cult., Botanical Garden Jena, Germany, Zimmermann 103/00 (BGJ), FN423746; E. mauritanica L., South Africa, Northern Cape, N of Pofadder, 29.02.2000, Zimmermann 12/00 (BGJ), FN423747 (specimen 2); E. meloformis Aiton, Ritz et al. (2003), AJ575343; E. meloformis fma. magna, Ritz et al. (2003), AJ575349; E. millotii Ursch & Leandri, ex cult., Botanical Garden Heidelberg, Germany, Rauh 8608 (HEID), FN423748 (specimen 2); E. monadenioides M.G. Gilbert, Ethiopia, Vlk 344 (BGJ), FN423749; E. neogossweileri Bruyns, Angola, Benguella, Lavranos s.n.; ex cult. International Succulent Institute (ISI 1261), leg. Forster 25311 (BRI), FN423750; E. nutans Lag., Canary Islands, Gran Canaria, Maspalomas, Faro II, 24.02.1996, Kuschel 646 (BGJ), FN423751; E. obesa Hook.f., Ritz et al. (2003), AJ575351; E.orthoclada Baker, Madagascar, Androy, 10.10.1998, Rauh 70626 (HEID), FN423752; E. pachypodioides Boiteau, Madagascar, Ankarana, 25.03.1991, Rauh 72072 (HEID), FN423753; E. parvicaruncula D.C. Hassall, Australia, Queensland, S of Boulia, 25.11.1999, Zimmermann 30/99 (BGJ), FN423754; E. pedilanthoides Denis, ex cult., Botanical Garden Heidelberg No. 142066, Germany, Rauh 66824 (HEID), FN423755; E. piscatoria Aiton, ex cult., Botanical Garden Berlin-Dahlem No. 075-01-96-10, Germany, (BGJ), FN423756; E. plumerioides Teijsm. ex Hassk., Australia, Queensland, Kennedy Hill Gorge, 21.06.1989, Forster 5402 (BRI), FN423757 (specimen 2); E. poeppigii (Klotzsch & Garcke) Boiss., Bolivia, La Paz, Nor Yungas, Cerro Uchimachi se Coroico, J. Müller 7040 (BGJ), FN423758;E. pseudoglobosa Marloth, Ritz et al. (2003), AJ575352 (specimen 1); AJ575353 (specimen 2); E. quartziticola Leandri, Madagascar, Ucal, Itremo, 24.10.1992, Rauh 73705 (HEID), FN423759 (specimen 2); E. sarcodes Boiss., Brazil, Bahia, Morro do Chapéu, 25.11.2000, Machado s.n. (BGJ), FN423760 (specimen 2); E. sarcostemmoides J.H. Willis, Australia, Queensland, W of Quilpie, 22.11.99, Zimmermann 23/99 (BGJ), FN423761; E. schizolepis F. Muell. ex Boiss., Australia, Western Australia, Packsaddle Plain, Kununurra, 04.03.1993, Mitchell 2851 (BRI), FN423762; E.schlechtendalii Boiss., ex cult., Botanical Garden Heidelberg, Germany, Rauh s.n. (BGJ), FN423763; E.scutiformis V.W. Steinm. & P.E. Berry, Cuba, Sancti Spíritus, Banao, 300-500m, 20.11.1975, Alvarez, Areces, Bisse, Dietrich & Lepper 28763 (BGJ), FN423764; E. sebsebei M.G. Gilbert, Ethiopia, Sidamo, Yavello, Vlk 326 (BGJ), FN423765; E. sipolisii N.E.Br., Brazil, Minas Gerais, Santa Barbara, Augusto de Lima, 06.08.1988, Eggli 1103 (ZSS), FN428581 (specimen 2); E. stellata Willd., South Africa, Eastern Cape, Motherwell, N of Port Elizabeth, 12.08.1997, Zimmermann 313/97 (BGJ), FN423766; E. sudanica A. Chev., ex cult., Botanical Garden Berlin-Dahlem No. 221-04-94-20, Germany (B), FN423767; E. susannae Marloth, Ritz et al. (2003), AJ575341 (specimen 1); AJ575340 (specimen 2); E. symmetrica A.C. White, R.A. Dyer & B. Sloane, Ritz et al. (2003); E. thinophila Phil., Chile, N of Huasco, 01.11.00, Ewest 03-11-00 (BGJ), FN423768 (specimen 2); E. trichadenia Pax, South Africa, Gauteng, Soutpan, NW of Pretoria, February 1998, Hardy s.n. (BGJ), FN423769; E.turbiniformis Chiov., Ritz et al. unpubl. AJ575319, E. valida N. E. Br., Ritz et al. (2003), AJ575350.
Neoguillauminiacleopatra (Baill.) Croizat New Caledonia, Mount Dzumac, 700m, 05.12.1980, Morat 6651 (BR), FN423770.
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Zimmermann, N.F.A., Ritz, C.M. & Hellwig, F.H. Further support for the phylogenetic relationships within Euphorbia L. (Euphorbiaceae) from nrITS and trnL–trnF IGS sequence data. Plant Syst Evol 286, 39–58 (2010). https://doi.org/10.1007/s00606-010-0272-7
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DOI: https://doi.org/10.1007/s00606-010-0272-7