Journal of Plant Research

, Volume 120, Issue 4, pp 511–522 | Cite as

Molecular phylogenetic analysis of Euphorbiaceae sensu stricto based on plastid and nuclear DNA sequences and ovule and seed character evolution

  • Toru TokuokaEmail author
Regular Paper


A phylogenetic analysis of Euphorbiaceae sensu stricto is presented using sequences from rbcL, atpB, matK and 18S rDNA from 85 species and 83 genera. The combined analysis of four molecular markers resulted in only one most parsimonious tree and also generated new supported clades, which include Euphorbioideae + Acalyphoideae s.s., subclades A2 + A3, subclades A5 + A6 and a clade uniting subclades A2–A8 within Acalyphoideae s.s. A palisadal exotegmen is a possible synapomorphy for all the Euphorbiaceae, except for the subfamily Peroideae. The presence of vascular bundles in the inner integument and a thick inner integument were shown to be synapomorphies for the clade of inaperturate and articulated crotonoids and for the large clade of Euphorbioideae, Acalyphoideae s.s., inaperturate and articulated crotonoids, respectively. Characters of the aril and vascular bundles in the outer integument are discussed. The selected embryological characters were seen to be highly correlated with the molecular phylogeny. When the results of molecular phylogenetic analysis of a previous study and this study were adjusted along with the selected embryological characters, all clades within Euphorbiaceae were supported except for a clade comprising Euphorbioideae + Acalyphoideae s.s. + inaperturate crotonoids + articulated crotonoids + Adenoclineae s.l. and a clade uniting subclades A4–A8 within Acalyphoideae s.s.


atpB Embryology Euphorbiaceae matK Phylogeny rbcL 18S rDNA 



I am grateful to Shawn Lum, Kenji Kaibe, Kipiro Damas and the curators of M.O., L and K.Y.O. for their assistance in obtaining materials used in the study; and to Hiroshi Tobe (K.Y.O.) and Louis Ronse De Craene (E) for helpful comments on the manuscript. The study was supported by a Grant-in-Aid for Scientific Research from the Japan Society for the Promotion of Science (17770069).

Supplementary material


  1. APG II (2003) An update of the angiosperm phylogeny group classification for the orders and families of flowering plants: APG II. Bot J Linn Soc 141:399–436Google Scholar
  2. Boesewinkel FD, Bouman F (1984) The seed: structure. In: Johri BM (ed) Embryology of angiosperms. Springer, Berlin, pp 567–610Google Scholar
  3. Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall MR, Price RA, Hills HG, Qiu Y-L, Kron KA, Pettig JH, Conti E, Palmer JD, Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD, Hedrén M, Gaut BS, Jansen RK, Kim K-J, Wimpee CF, Smith JF, Furnier GR, Strauss SH, Xiang QY, Plunkett GM, Soltis PS, Swensen SM, Williams SE, Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn GH Jr, Graham SW, Barrett SCH, Dayanandan S, Albert VA (1993) Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Ann Missouri Bot Gard 80:528–580CrossRefGoogle Scholar
  4. Chase MW, Zmarzty S, Lledó MD, Wurdack KJ, Swensen SM, Fay MF (2002) When in doubt, put it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bull 57:141–181Google Scholar
  5. Davis CC, Chase MW (2004) Elatinaceae are sister to Malpighiaceae; Peridiscaceae belong to Saxifragales. Am J Bot 91(2):262–273Google Scholar
  6. Davis CC, Wurdack KJ (2004) Host-to-parasite gene transfer in flowering plants: Phylogenetic evidence from Malpighiales. Science 305(5684):676–678PubMedCrossRefGoogle Scholar
  7. Davis CC, Webb CO, Wurdack KJ, Jaramillo CA, Donoghue MJ (2005) Explosive radiation of Malpighiales supports a Mid-Cretaceous origin of modern tropical rain forests. Am Nat 165:E36–E65PubMedCrossRefGoogle Scholar
  8. Felsenstein J (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39:783–791CrossRefGoogle Scholar
  9. Govaerts R, Frodin DG, Radcliffe-Smith A (2000) World checklist and bibliography of Euphorbiaceae (with Pandaceae). Redwood, Trowbridge WiltshireGoogle Scholar
  10. Hilu KW, Borsch T, Müller K, Soltis DE, Soltis PS, Savolainen V, Chase MW, Powell MP, Alice LA, Evans R, Sauquet H, Neinhuis C, Slotta TAB, Rohwer JG, Campbell CS, Chatrou LW (2003) Angiosperm phylogeny based on matK sequence information. Am J Bot 90(2):1758–1776Google Scholar
  11. Mason-Gamer RJ, Kellogg EA (1996) Testing for phylogenetic conflict among molecular data sets in the tribe Triticeae (Gramineae). Syst Biol 45(4):524–545CrossRefGoogle Scholar
  12. Savolainen V, Fay MF, Albach DC, Backlund A, van der Bank M, Cameron KM, Johnson SA, Lledó MD, Pintaud JC, Powell M, Sheahan MC, Soltis DE, Soltis PS, Weston P, Whitten WM, Wurdack KJ, Chase MW (2000a) Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bull 55:257–309Google Scholar
  13. Savolainen V, Chase MW, Hoot SB, Morton CM, Soltis DE, Bayer C, Fay MF, de Bruijn AY, Sullivan S, Qiu YL (2000b) Phylogenetics of flowering plants based on combined analysis of plastid atpB and rbcL gene sequences. Syst Biol 49:306–362PubMedCrossRefGoogle Scholar
  14. Schmid R (1986) On cornerian and other terminology of angiospermous and gymnospermous seed coats: historical perspective and terminological recommendations. Taxon 35:476–491CrossRefGoogle Scholar
  15. Soltis DE, Soltis PS, Nickrent DL, Johnson LA, Hahn WJ, Hoot SB, Sweere JA, Kuzoff RK, Kron KA, Chase MW, Swensen SM, Zimmer EA, Chaw S-M, Gillespie LJ, Kress WJ, Sytsma KJ (1997) Angiosperm phylogeny inferred from 18S ribosomal DNA sequences. Ann Missouri Bot Gard 84:1–49CrossRefGoogle Scholar
  16. Soltis DE, Soltis PS, Chase MW, Mort ME, Albach DC, Zanis M, Savolainen V, Hahn WH, Hoot SB, Fay MF, Axtell M, Swensen SM, Prince LM, Kress WJ, Nixon KC, Farris JS (2000) Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Bot J Linn Soc 133:381–461CrossRefGoogle Scholar
  17. Swofford DL (2001) PAUP: Phylogenetic analysis using parsimony, ver 4.0b10. Sinauer, SunderlandGoogle Scholar
  18. Tokuoka T, Tobe H (1995) Embryology and systematics of Euphorbiaceae sens. lat.: a review and perspective. J Plant Res 108:97–96CrossRefGoogle Scholar
  19. Tokuoka T, Tobe H (1998) Ovules and seeds in Crotonoideae (Euphorbiaceae): structure and systematic implications. Bot Jahrb Syst 120:165–186Google Scholar
  20. Tokuoka T, Tobe H (1999) Embryology of tribe Drypeteae, an enigmatic taxon of Euphorbiaceae. Pl Syst Evol 215:189–208CrossRefGoogle Scholar
  21. Tokuoka T, Tobe H (2001) Ovules and seeds in subfamily Phyllanthoideae (Euphorbiaceae): structure and systematic implications. J Plant Res 114:75–92CrossRefGoogle Scholar
  22. Tokuoka T, Tobe H (2002) Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications. J Plant Res 115:361–374PubMedCrossRefGoogle Scholar
  23. Tokuoka T, Tobe H (2003) Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications. J Plant Res 116:355–380PubMedCrossRefGoogle Scholar
  24. Tokuoka T, Tobe H (2006) Phylogenetic analyses of Malpighiales using plastid and nuclear DNA sequences, with particular reference to the embryology of Euphorbiaceae sens. str. J Plant Res (in press)Google Scholar
  25. Webster GL (1987) The saga of the spurges: a review of classification and relationships in the Euphorbiales. Bot J Linn Soc 94:3–46Google Scholar
  26. Webster GL (1994a) Classification of the Euphorbiaceae. Ann Missouri Bot Gard 81:3–32CrossRefGoogle Scholar
  27. Webster GL (1994b) Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Ann Missouri Bot Gard 81:33–144CrossRefGoogle Scholar
  28. Wurdack KJ (2002) The molecular systematics and evolution of Euphorbiaceae sensu lato. PhD dissertation, University of North CarolinaGoogle Scholar
  29. Wurdack KJ, Hoffman P, Chase MW (2005) Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F sequences. Am J Bot 92:1397–1420Google Scholar

Copyright information

© The Botanical Society of Japan and Springer 2007

Authors and Affiliations

  1. 1.Department of Natural Environmental Sciences, Faculty of Integrated Human StudiesKyoto UniversityKyotoJapan

Personalised recommendations