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Home , Allantoma, Asteranthos, Brazzeia, Careya, Cariniana, Chydenanthus

TAXON 47 - NOVEMBER 1998 817

Recircumscription of the

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 . 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 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 and (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 nut, the edible seed of Bertholletia excelsa Humb. &. BonpJ., and the widely cultivated cannonball , 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 ". 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 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 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. . 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 , 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, Pierre ex Engl. and Baill. He also described subfamily Rhaptopetaloideae O. Appel to accommodate the remaining genera of the Scytopetalaceae ( Oliv., Pierrina Engl., and 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 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. amoreaRoxb. Chase2256, K AF077655 AF077640 Carinlanalegalis(Mart.)Kuntze Kew 1991-169 Z80179 Carinlanaestrellensis (Raddi) Kew 1991·170 AF077647 Kuntze excalsa (Blume) BogorV.A.S. Z80180 AF077641 Miers 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 poeppigiana O. Berg Chase 330, K Z80175 AF077636 prancei S. A. Morl Ducke Res. 104, INPA AF077651 AF077630 Hook. & Chase 329, K Z80173 AF077649 Planch. afrlcanus Merr. Morl & Kisseadoo 19241, AF077654 AF077638 NY Planchonla valida Blume Bogor XII.B.VII Z80176 AF077642

Primulales Myrsinaceae Rapanea fe"uginea Mez Chase 334, K Z80204 Maesa myrsinoldes Lev. Kew 1992-3180 Z80203 PniTJulaceae Androsace spinulifera Knuth Chase 954, K Z80205 Theophrastaceae Jacquinia umbellata A. DC. Axelrod 4552, UPR Z80206

Rosales Byblidaceae gorgonias Planch. Goldblatt 5393, MO L01950 Hydrangeaceae Hydrangea macrophylla Torr. Morgan 2150, WS L11187

Sa"aceniales • ..•. :> Sa"aceniaceae Oarlingtonia califomica Torr. & Williams 0101, LVC L02432 A. Gray Sa"acenia flava L. Chase 144, NCBG L01952 .,1(" :~-. Theales chinensis Planch. Kron2117, NCU L01882 Clems toclethra lasioclada Edinburgh 1909-1001 Z80172 Maxim. Saurauia nepaulensis DC. Chase 1096, K Z83147 AF080486 Norantea guianensis Aubl. Mori 22996, NY Z80200 Scytopetalaceae Brazzeia sp. ' Brete/er& al.12916, WAG AF077656 AF077644 Oubanguia alata Baker f. Gereau & al. 5202, MO Z80201 Oubangula sp. Breteler & al.13096, WAG za0202 AF077643 Theaceae japonica Thunb. Kew 1992·1192 Z80207 Schims supetba Gardiler & Lammers, F Z80208 Champ. Stuartia pseudocamellia Maxim. Kew 1949-66901 Z80209 Temstroemla stahliiKrug & Axelrod 4538, UPR Z80211 Urban TetramerislaCBBe Tetramerlsta sp. Coode 7925, K Z80199

Violales FouquieriaCBBe FouquierlasplendBns Engelm. Matthae/, BG 860162 L11675 Idria columnaria Kellogg UCI Arbor. Z80210

DNA extraction. - Silica-dried and herbarium material, "0.2-0.3 g", were ground into a fine paste and incubated according to the shortened 2X CTAB proce­ dure of Doyle & Doyle (1987). Proteins were removed with SEVAG (24 : 1 chloro­ form: isoamyl alcohol), followed by a final purification using equilibrium density­

dependent centrifugation in CsCl2-ethidium bromide (1.55 g/ml). Gene amplification and sequencing. - Amplification for the rbcL gene was per­ formed using the following oligonucleotide primers described by Olmstead & al. (1992) and for the trnL gene using the primers described by Taberlet & al. (1991). L. and Brazzeia BaHI. rbcL sequences are based on partial sequences due to amplification problems. Slight modifications to the reaction conditions were found

to be necessary for some taxa, e.g., the MgCl2, BSA, and DMSO concentrations TAXON 47 - NOVEMBER 1998 821

varied. Amplified products were cleaned using Promega Magic mini-columns fol­ lowing protocols provided by the manufacturer. Cleaned products were then directly cycle sequenced using the AmpliTaq DNA polymerase cycle sequencing ready re­ action kit (Perkin-Elmer Corp.) on a pre-programmed Perkin-Elmer thermal cycler. Samples were then cleaned and run on a ABI 373A DNA sequencer, and data were collected on a Macintosh platform. Raw sequencing data were edited using the pro­ gram Sequencher (Gene Codes Corporation). Visual alignment was then accom­ plished with varying degrees of ease; the conserved nature of the rbcL made it a fairly simple task; however, the number and occurrence of indels within trnL made its alignment more difficult (exclusion of bp 191-245 and 448-458). Analysis of data. - All data sets were analysed using the parsimony algorithm software package PAUP 3.1.1 (Swofford, 1993). To examine molecular support for subfamilial relationships and to clarify relationships of Lecythidaceae among other families and orders we examined 47 rbcL sequences of 25 families (sensu lato: Cronquist, 1981). All analyses support recognition of the monophyly of the "asterid III" grouping of Chase & al. (1993). From these results we selected the restricted data set. This data set was analysed using the heuristic search option set for Tree Bisection Reconnection (TBR), MULPARS, and Steepest Descent, with all characters weighted equally and unordered (Fitch parsimony: Fitch, 1971). All trees were swapped to completion. Chrysophyllum L. of Sapotaceae Juss. and Saurauia Willd. of Actinidiaceae Hutch. were used as the outgroups, based on the results from the larger analyses. Branch lengths for trees were calculated using ACCTRAN (acceler­ ated transformation optimisation) option in PAUP. Relative support for clades identi­ fied by parsimony analysis was assessed using jackknifing (JK; 100 replicates) and bootstrapping (BS; 100 replicates) indices (PAUPstar version 4.0.0d64; Swofford, unpublished).

Results Higher level analysis. - The higher level rbcL heuristic analysis using 47 se­ quences produced five trees of 1144 steps, CI = 0.50 and RI = 0.53 (Morton & al. 1997). Lecythidaceae are paraphyletic with Scytopetalaceae excluded, and the Lecy­ thidaceaelScytopetalaceae clade is well supported by jackknifing and bootstrapping indices (fig. not shown). Relationships indicated by combined rbcL, trnL, and morphological data set. ­ The analysis of the combined molecular data sets and the morphological data set results in three trees of 471 steps with CI =0.71 and RI =0.70. In all trees Lecythi­ daceae are paraphyletic and all major branches are supported (Fig 1.) These trees support the monophyly of three of the four subfamilies (Planchonioideae, Lecythi­ doideae, and Foetidioideae) recognised by Prance & Mori (1979), with the fourth, Napoleonaeoideae, forming two separate clades. In the first major clade Planchonioideae ( J. R. Forst. & G. Forst., Careya Roxb., Chydenanthus Miers, Petersianthus Merr., and Blume) are monophyletic and sister to Comm. ex Lam., the only of Foetidi­ oideae. The second major clade contains Lecythidoideae ( Miers, Berthol­

) letia Humb. & Bonpl., Casar., Corythophora Knuth, Aubl., i Couroupita Aubl., Mart., Grias, Gustavia L., and Lecythis Loefl.); this s subfamily is supported, although relationships among the genera lack support. Aster­ ------v-­

822 TAXON 47 - NOVEMBER 1998

anthos(Napoleonaeoideae), Oubanguia and Brazzeia (Scytopetalaceae) form a strongly supported clade in this analysis. This clade is sister to Planchonioideae, Foetidioideae

Lecyth/s

Eschwe/ltlra

Corythophora

Allantoma

Couratarl ythidoideae Cllrin/ana

Ekn1hoIIetIa

Grlas

Courouplta

GIAltlwIa 62162 Foetld/a Foetidioldeae

Peters/anthus 10Qf1oo

Ianchonioideae Planchon/a 10qJ98 Barrlngtonla

Chydenanthus 30 1 100 :::'UM}ytopetalOideae

'----&-- Asteranthos

Napoleonaea Napoleonaeoideae

.....:IiL-______Outgroups L... -1Saurau/a

Fig. I. One of three most parsimonious trees, using rbeL sequence, tmL sequence, anatomical, cytological. and morphological characters, resulting from a heuristic search. Sapotaceae and Aeti­ nidiaeeae served as outgroups. Length =471 steps, CI =0.71, RI =0.70. Segment lengths (ACCTRAN optimisation) are indicated above the branches, bootstrappingljackknifing indices are below each branch. TAXON47-NOVEMBERIW8 823

and Lecythidoideae. Napoleonaea is basal to the rest of the family. The only other differences found among the three trees are the relative positions of genera within the Lecythidoideae.

Taxonomy Relationships of Lecythidaceae. - Several large analyses, using rbcL, atpB and 18S, strongly support the monophyly of the "asterid IIf' grouping (Morton & aI., 1997; Savolainen & aI., 1996; Soltis & at, 1996; Morton & a1. 1996; Chase & aI., ·1993; Olmstead & aI., 1992, 1993). Resolution is poor in many instances in the higher level rbcL analysis, and branch support for specific groupings is weak. How­ ever, Sapotaceae form a sister clade with LecythidaceaelScytopetalaceae in all trees produced (Morton & aI., 1997). The phylogenetic position of Lecythidaceae, does not support a relationship with Myrtales Rchb. (Hutchinson, 1973; Takhtajan, 1980), Malvales Dumort. (Cronquist, 1957), or Urticales Dumort. (Stebbins, 1974). Furthermore these analyses do not support a relationship of Scytopetalaceae with Olacaceae Mirbel ex DC. (Oliver, 1868), Malvales (Takhtajan, 1980), Tiliales (Hutchinson, 1973), or Ochnales Hutch. ex Reveal (Takhtajan, 1987). This analysis does support the past alignment of Le­ cythidaceae and Scytopetalaceae with a loosely defined Theales Lindl. (Cronquist, 1981;' Dahlgren, 1983; Thome, 1992) and with 'some members of Violales Perleb (Stebbins, 1974). We acknowledge that more morphological and molecular data are needed to provide additional interfamilial resolution. Monophyly ojLecythidaceae/Scytopetalaceae. - Both families share a number of morphological features, such as: alternate, simple leaves, perfect flowers, cortical bundles in stem, numerous stamens, three-aperturate pollen, axile placentation, and a bitegmic-tenuinucellate ovule. While most of these characters occur widely through­ out the dicots, therefore having limited utility in establishing phylogenetic relation­ ships by themselves, two cha'taeteristic features, viz., bitegmic-tenuinucellate (B-1) ovules and cortical bundles in stems, do have greater taxonomic value at the family level because of their restricted occurrence (Davis, 1966; Philipson, 1974; Cron­ quist, 1981; Metcalfe & Chalk, 1983). Within the combined data sets three morpho­ logical synapomorphies were found (viz., numerous stamens, bitegmic ovules, and cortical bundles in the stem), suggesting a close phylogenetic relationship between the two families. As a result we have concluded that the Lecythidaceae should be divided into five subfamilies which we describe below. The Scytopetalaceae are formally included as part of the Lecythidaceae for the first time. Planchonioideae Nied, in Engler & Prantl, Nat. Pflanzenfam. 3(7): 30-33. 1892. ­ Type: Planchonia Blume. Flowers actinomorphic; present, corona of staminal origin absent, inner or outer whorl of stamens sometimes staminodial, the base of the stamens fused into a staminal ring; intrastaminal disk present; pollen syntricolpate with a specialised marginal ridge; ovary 2-4-locular, placenta columnar. Fruit indehiscent. Secondary xylem with cortical bundles reversely oriented with the xylem on the outside. x = 13. The Planchonioideae consist of six genera, all palaeotropical in distribution. All except Barringtonia, the largest genus with 41 species, have a limited number of species ranging from two to eight species (Abdulmajidia Whitm., 2; Petersianthus, 2; 824 TAXON 47 - NOVEMBER 1998 and Chydenanthus, 2; Careya, 4; Planchonia, 8). All of the genera have a limited geographic range except Barringtonia, which is widespread from E. Africa through Madagascar to tropical Asia. Tsou's (1994a) embryological study found the floral morphology of all six genera to be similar, containing a number of easily recognis­ able but common characters to the family. In our cladistic examination of five of the six genera, we found that the monophyly of Planchonioideae is supported by two unambiguous morphological synapomorphies: syntricolpate pollen, and pollen with a thickened colpus ridge. The pollen of Planchonioideae is unique among angio­ sperms in the combined presence of syncolporate apertures and a thickened extexi­ nous ridge along the colpus margin (Erdtman, 1971; Muller, 1972; Tsou, 1994 a, b). All other members of the family possess tricolp(or)ate pollen.

Foetidioideae Nied. in Engler & Prantl, Nat. Pflanzenfam. 3(7): 29-30. 1892 == [tri­ bus] Foetidieae Knuth in Engler, Pflanzenr. 105: 62-65. 1939 == [family] Foetidi­ aceae (Nied.) Airy Shaw in Kew Bull. 18: 258. 1965. - Type: Foetidia Comm. ex Lam. Flowers actinomorphic; petals absent, corona of stamlnalnrigin absent, stamino­ des absent, the base of the stamens weakly fused and not forming a staminal ring; intrastaminal disk present; pollen tricolpate; ovary 4-10cular, placenta peltate. Fruit indehiscent. Secondary xylem with cortical bundles reversely.oriented. Chromosome number unknown. This monogeneric subfamily consists of 17 species, all of which are confined to Madagascar except for one found in Mauritius and another in E. Africa. Tsou (1994a) stated: "the embryological features of Foetidia match quite well with those of Planchonioideae and Lecythidoideae, differing most prominently in its well­ developed integumentary fusion". The Planchonioideae-Foetidioideae clade is supported by one morphological synapomorphy, viz., cortical bundles in the stem with reverse orientation. The Foe- . tidioideae have four morphological autapomorphies: apetalous flowers, introrse anther dehiscence, nearly free filaments, and the presence of silica; all of these char­ acter states are paralleled in other genera of Lecythidaceae.

Lecythidoideae Nied. in Engler & Prantl, Nat. Pflanzenfam. 3(7): 34-41. 1892 == [tribus] Lecythideae Bercht. & J. Presl, Pfir, Rostlin 2: 378. 1825 == [family] Le­ cythidaceae Poit. in Mem. Mus. Hist. Nat. 13: 143. 1825, nom. cons. - Type: Le­ cythis Loefl. Flowers actinomorphic or zygomorphic; petals present, corona of staminal origin absent, stamens arising from a connate staminal ring in Gustavia, Grias, and Allan­ toma, the staminal ring slightly expanded on one side in Cariniana, and markedly expanded into a strap-like ligule with an enlarged hood at the apex in the remaining genera; intrastaminal disk absent; pollen tricolp(or)ate; ovary usually 2-, 4-, or 6­ locular, placenta axile or modified axile. dehiscent or indehiscent. Secondary xylem with cortical bundles normally oriented. x = 17. The Lecythidoideae are composed of ten genera endemic to the Neotropics. In the combined analysis using all ten genera, the·Lecythidoideae are defined by a single non-molecular synapomorphy, viz., a basic chromosome number of x =17. TAXON 47 - NOVEMBER 1998 825

Scytopetaloideae O. Appel in Bot. J. Linn. Soc. 121: 225. 1996 == [family] Scytope­ talaceae Engl. in Engler & Prantl, Nat. Pflanzenfam., Nachtr. 1: 242. 1897, nom cons.- Type: Scytopetalum Pierre ex Engl. Flowers actinomorphic; petals absent or present with a corona of staminal origin, staminodes absent, the base of the stamens fused but not forming a staminal ring; intrastaminal disk mostly absent; pollen tricolp(orio)ate; ovary 3- to 8-locular, pla­ centa axile. Fruit indehiscent except for Oubanguia. Secondary xylem with normally oriented coritcal bundles. x = 11, 18. Scytopetaloideae, as here circumscribed, are comprised of Asteranthos, Brazzeia, and Oubanguia. All except Asteranthos are restricted to tropical W. Africa. Aster­ anthos is confined to the upper Rio Negro in Colombia, Venezuela, and Brazil. Based on these results, Pierrina Engl., Rhaptopetalum Oliv., and of course-Ssyro­ petalum, should probably be considered as members of Scytopetaloideae as well, but no material was available for molecular study and therefore their placement is tenta­ tive until additional examinations are completed. The close association between Asteranthos and Scytopetalaceae is supported by the data of Tsou (1994a) and Appel (1996). Our analysis found one unambiguous morphological synapomorphy for the gen­ era of Scytopetaloideae: the presence of ruminate endosperm. Ruminate endosperm is found in all Scytopetaloideae except Oubanguia, but this genus occasionally is reported also to possess the feature (Letouzey, 1960). In contrast, all other Lecythi­ daceae lack endosperm.

Napoleonaeoideae Nied. in Engler & Prantl, Nat. Pflanzenfam. 3(7): 33-34. 1892; == [family] Napoleonaeaceae A. Rich. in Bory, Diet, Class. Hist. Nat. 11: 432. 1827 == [subtribus] Napoleonaeinaeae Benth. & Hook. f., Gen. PI. 1: 723-727. 1865 == [tribus] Napoleonaeeae Knuth in Engler, Pflanzenr. 105: 67-73. 1939. ­ Type: Napoleonaea P. Beauv., ....~, ..~~ ... Flowers actinomorphic; petals absent, corona of staminal origin present, stamino­ des present in Napoleonaea, absent in Crateranthus, the base of the stamens fused into a staminal ring; intrastaminal disk present in Napoleonaea, absent in Crateran­ thus; pollen tricolpate; ovary 3- or 5-locular, placenta columnar. Fruit indehiscent. Secondary xylem with cortical bundles normally oriented with phloem on the out­ side. x = 16 in Napoleonaea, unknown in Crateranthus. Based on morphological data Napoleonaeoideae consist of Napoleonaea (8 sp.) and Crateranthus Baker f. (3 sp.), both of W. African distribution. This circum­ scription is consistent with Niedenzu's initial description which included Napoleo­ naea and Crateranthus in the subfamily while he placed Asteranthos in its own family. No material of Crateranthus was available for DNA study, and, therefore we are unable to comment further on its relationships using molecular data. Tsou (1994a) stated that "if most embryological features of Asteranthos are considered to be plesiomorphies of the Lecythidaceae s.l., then Crateranthus and Napoleonaea probably represent, in terms of embryology, a distinct clade branching from the base of the group". Our analysis based on non-molecular data confirms Tsou's placement ,of these taxa. Napoleonaeoideae are characterised by four non-molecular autapomorphies: the presence of a staminal corona, extrorse anther dehiscence, a basic chromosome -~...... ------~.a;.;; •.-­ I

826 TAXON 47 - NOVEMBER 1998

number of x =16, and apetalous flowers. The presence of a staminal corona and apetalous flowers are homoplastic within Lecythidaceae s.l. Relationships of Lecythidaceae and Sapotaceae. - Molecular data from several higher level analyses, using rbcL and atpB data (Morton & al., 1997; Savolainen & al., 1996; Morton & al. 1996; Chase & at, 1993; Olmstead & al., 1992, 1993), sug­ gest that the Sapotaceae and Lecythidaceae are sister families. We acknowledge that additional sampling from the Sapotaceae using non-molecular data is needed to increase our understanding of evolutionary patterns within these groups.

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