TAXON 47 - NOVEMBER 1998 817 Recircumscription of the Lecythidaceae Cynthia M. Morton'", Ghillean T. Prance', Scott A. Mori4 & Lucy G. Thorburn' Summary Morton. C. M.• Prance, G. T., Mori, S. A. & Thorburn. L. G.: Recircumscriplion of the Le­ cythidaceae. ­ Taxon 47: 817-827. 1998. -ISSN 004Q-0262. The phylogenetic relationships of the genera of Lecythidaceae and representatives of Scyto­ petalaceae were assessed using cladistic analysis of both molecular (rbcL and trnL se­ quences) and morphological data. The results show that the pantropical family Lecythida­ ceae is paraphyletic. Support was found for the monophyly of three of the four subfamilies: Lecythidoideae, Planchonioideae, and Foetidioideae. The fourth subfamily, Napoleonaeol­ deae, was found to be paraphyletic, with members of the Scytopetalaceae being nested within it forming a strong clade with Asteranthos. Both families share a number of mor­ phological features, including several distinct characters such as cortical bundles in the stem. The combined analysis produced three trees of 471 steps and consistency index Cl = 0.71 and retention index Rl = 0.70. Asteranthos !'P.~, members of Scytopetalaceae should be treated as a subfamily of Lecythidaceae, while Napoleonaea and Crateranthus (the latter based solely on morphological features) should remain in the subfamily Napoleo­ naeoideae.The Lecythldaceaeare recircumscribed, and Asteranthosand members of Scyto­ peta/aceae are included in Scytopetaloideae. A formal·llWJ!J-pmic synopsis accommodating this new circumscription is presented. Introduction The Lecythidaceae Poit, are 8 pantropical family of trees and shrubs consisting of . 20 genera split into four subfamilies in contemporary classifications (Cronquist, 1981; Prance & Mori, 1979; Takhtajan, 1987; ~ri & Prance, 1990; Thome, 1992). It is best known by the Brazil nut, the edible seed of Bertholletia excelsa Humb. &. BonpJ., and the widely cultivated cannonball tree, Couroupita guianensis Aubl. Taxonomic alignments for the family have varied during the last century. Poiteau (1825) stated that Lecythidaceae should be treated as a distinct family, separate from Myrtaceae Juss., a move supported by Lindley (1846) on the evidence of the "great almond-like seeds and alternate, often serrated, non-punctate leaves". He also placed Napoleonaea P. Beauv. and Asteranthos Desf. within Belvisiaceae R. Br. near the Barringtoniaceae F. Rudolphi. The first monograph was published by Miers (1874) who placed great emphasis on fruit characters, especially minute differences in shape and form. His species were based on a morphological rather than the genetic concept applied by modem taxonomists, however, his monograph provided accurate, well-illustrated accounts of species representing all genera known at the time (Prance & Mori, 1979). Miers's families Lecythidaceae and Barringtoniaceae are equivalent to the subfamilies Le­ cythidoideae Nied. and Planchonioideae Nied. of Niedenzu (1892) whose treatment has been followed by most modem workers (e.g., Prance & Mori, 1979). I Uni versity of Reading, School of Plant Sciences, Whiteknights, PO Box 221, Reading RG6 2AS, U.K. 1 Auburn University, Department of Botany and Microbiology. Auburn. AL 36849-5407, U.S.A. l Royal Botanic Gardens. Kew, Richmond. Surrey, TW9 3AB, U.K. i'l l' • New York Botanical Garden. Bronx, NY 10458-5126, U.S.A. 818 TAXON 47 - NOVEMBER 1998 The second monographer, Knuth (1939), added little to the understanding of the family. According to Prance & Mori (1979) he was a "splitter", having extremely limited material, and hence produced a classification independent of the biology of the family. In 1945, Pichon proposed a new classification, dividing the family into 3 subfamilies and 12 tribes. He provided information about floral, seed, and embryo­ logical characteristics of the taxa. The most recent monograph of Prance & Mori (1979) and Mori & Prance (1990), treated the Neotropical species in detail and summarised the Old World groups. Their circumscription of the family recognised 20 genera grouped into four subfamilies: Planchonioideae, Lecythidoideae, Foetidioideae Nied., and Napoleonaeoideae Nied. Tsou's embryological study of the family (1994a) was the first comprehensive study of this kind. She stated that the subfamilies Lecythidoideae and Planchoni­ oideae form the core-group of Lecythidaceae. In contrast, she argued that Foetidi­ oideae and Napoleonaeoideae should be recognised as separate families, although she did not propose a formal classification. The close affinities between the non core Lecythidaceae and Scytopetalaceae Engl. were also noted. The work of Morton & al. (1997), using rbcL sequences, morphological, ana­ tomical, and cytological data, suggested that Lecythidaceae .. ~,,_ circumscribed by Prance & Mori (1979) are paraphyletic. Moreover, Tsou's promotion of the non-core Lecythidaceae as separate families (Tsou, 1994a) lacked support. Morton & al. (1997) also found, using both molecular and morphological combmed data sets in a cladistic analysis, that Asteranthosgrouped with Scytopetalaceae. Scytopetalaceae are a family of five genera with c. 22 species consisting of trees, shrubs, and lianas restricted to W. Africa. The family is defined by the same features as found in Lecythidaceae, except that it can further be characterised by having ex­ stipulate leaves, anisocytic stomata" hypogynous flowers, and an anatropous ovule (Appel, 1996; Cronquist, 1981; Letouzey, 1960, 1978). Scytopetalaceae have been included in four different orders: Ochnales (Takhtajan, 1987), Malvales (Takhtajan, 1980), Theales (Dahlgren, 1983; Cronquist, 1981; Thome, 1992), and Tiliales (Hut­ chinson, 1973). The most recent discussion of Scytopetalaceae has been provided by Appel (1996). Appel supported Tsou's core of Lecythidaceae, but recommended the re­ moval of Asteranthosfrom Napoleonaeoideae, its inclusion in Scytopetalaceae, and the formation of a' subfamily,Scytopetaloideae O. Appel within Scytopetalaceae, to include Asteranthos, Scytopetalum Pierre ex Engl. and Oubanguia Baill. He also described subfamily Rhaptopetaloideae O. Appel to accommodate the remaining genera of the Scytopetalaceae (Rhaptopetalum Oliv., Pierrina Engl., and Brazzeia Baill.). Apple's inclusion of Asteranthosin the Scytopetalaceae is based on the pre­ sumption that the pseudocorolla of both are staminal in origin, as well as on floral morphological and seed morphological characteristics (e.g., the co-occurence of ruminate endosperm in Asteranthosand other members of his Scytopetaloideae). Recent molecular examinations supports the inclusion of Scytopetalaceae within Lecythidaceae rather than the placement of Asteranthos in a segregate Scytopeta­ laceae as proposed by both Tsou (1994a) and Appel (1996). Materials and methods Plant material. - A list of specimens examined, voucher depositions, and gene- ( bank accession numbers are listed in Table 1. TAXON47 - NOVEMBER 1998 819 Table 1. Species analysed in mel. and trnL sequence analysis with voucher information arid DNA database accession numbers. Taxa are arranged alphabetically by order,then family and species. Order Species Voucher/source mel database tmLdatabase Family '" accessionNo. accession No. ,Coma/es J Nyssaceae Nyssa ogeehe Marsh. US NatlArbor. L11228 Comaceae AueubajaponicaThunb. US NatlArbor. L11210 'Diapensiales Diapensiaeeae Galax aphyllaL. Kron, NCU Z80184 Dillenia/es Dilleniaeese Dillenlaindica L. Chase 234,NCO. L01903 " :'>1:" £bena/es Ebenaeeae DiospyroskakiL. Kew 1992·3634 Z80185 Euelesnata/ensis A. DC. Goldblan 9275,K Z80186 SapQtaceae ArganiaspinosaSkeels Kew1988-1 898 Z80187 Chrysophyllum oliviforme L. Chase 127, NCU L12607 AF077650 Pouterlaeerwah(F. M. Bailey) Kew 1986-2962 Z80188 BaehnJ Styraeaeeae HalaslacarolinaL. Kew 1982-4193 Z80190 Rehderodendron maerocarpum Kew 1969-32516 Z80191 Hu I StyraxameriCanusLam. Kron3002, NCU L12623 StyraxjaponieusSieb.& Zucco Kew1985-8633 Ztlo189 Symplocaeeae Symploeos costataCholsyex BogorXVlfI.B4Clbodas Z80192 Zoll. Symploeos ferruginea Roxb. BogorXVIII.B6Clbodas Z80193 Ericales Ericaceae Craiblodendron yunnanense Kew1982-5379 Z80195 W.W.Sm. Enkianthuscampanulatus ArnoldArbor. 14528-C L12616 G. Nicholson Rhodothamnus ehaml!Jlf/stus Kew 1989-459 Z80194 Rchb. .0<'- • Cyrillaeeae CyrillaraeemiRora L. Kron, NCU L01900 C/ethraeeae ClethraamoreaSol. Kew 1987-4005 Z80212 Gerania/es Balsaminaceae ImpatiensgordoniiHomeex Chase 563,K Z80196 Baker Impatiens repensMoon Kew 1969-51717 Z80171 Lecythida/es Lecythidacaae AllantomallneataMiers DuckeRes. 119,INPA AF077657 AF077645 AsteranthosbrasilIensIs Des!. Mori& al. 21856,NY Z80198 AF077648 Barrlngtonla asIaticaKurz Chase328,K Z80174 AF077639 Bertholletlaexeelsa Humb.& Chase 326,K Z80178 AF077635 Bonpl. Careya amoreaRoxb. Chase2256, K AF077655 AF077640 Carinlanalegalis(Mart.)Kuntze Kew 1991-169 Z80179 Carinlanaestrellensis (Raddi) Kew 1991·170 AF077647 Kuntze Chydenanthus excalsa (Blume) BogorV.A.S. Z80180 AF077641 Miers Corythophora rimosa DuckeRes. 164,INPA AF071653 AF071633 W. A. Rodrigues CourataritauariO. Berg DuckeRes.2595.INPA Z80171 AF077646 Couroupitagulanensis AubJ. Chase 327,K Z80181 AF071632 Eschweilara odora(Poepp.) Miers Granville 5086(CAY) Z80182 AF071634 FoetidiaasymetricaH. Perrier Chase 1446, K Z80183 AF071637 " . ,', 820 TAXON 47 - NOVEMBER 1998 Table 1. (continued). Order Species Voucherlsource tbeL database tml database Family accession No. accession No. Grias cauliflora L. Chase 2196, K AF077652 AF077631 Gustavia poeppigiana O. Berg Chase 330, K Z80175 AF077636