Amborellaceae - Species Plantarum Programme

AMBORELLACEAE
Flora of the World
14. Amborellaceae

Cover illustration:
Amborella trichopoda Baill. Reproduced with permission, from Fl. Nouvelle-Calédonie &
Dépend. 11: 159 (1982). Drawn by H. Lamourdedieu.

SPECIES PLANTARUM
FLORA OF THE WORLD
PART 14. AMBORELLACEAE
by Joël Jérémie, Porter P. Lowry II & Frédéric Tronchet
Published for the Species Plantarum Programme
by the Conservatoire et Jardin botaniques de la Ville de Genève

Editions des Conservatoire
et Jardin botaniques
EDITORS
Receiving editor: J. Kirschner
Editors: C. E. Anderson, K. Marhold
EXECUTIVE COUNCIL OF THE SPECIES PLANTARUM PROGRAMME
G. F. Smith, National Botanical Institute, Pretoria, South Africa (Chairman)
J. Kirschner, Inst. Bot., Pr ° uhonice, Czech Republic (Vice-Chairman, Receiving Editor)
K. Marhold, Slovak Academy of Sciences, Bratislava, Slovakia (Editor)
W. Thomas, New York Botanic Gardens, Bronx, New York, USA (Secretary)
W. L. Wagner, Smithonian Institution, Washington DC, USA (Treasurer)
P.-A. Loizeau, Conservatoire et Jardin botaniques, Genève, Switzerland (Publisher)
The work may be cited as:
Joël Jérémie, Porter P. Lowry II & Frédéric Tronchet, Amborellaceae,
Species Plantarum: Flora of the World, Part 14: 1-7 (2008)
The book is available from:
Conservatoire et Jardin botaniques de la Ville de Genève
Case postale 60
CH-1292 Chambésy
editions.cjb@ville-ge.ch
http://www.ville-ge.ch/cjb/
ISBN 978-2-8277-0453-8

CONTENTS
Contributors.............................................................................................................................. iv
Species Plantarum Committee................................................................................................... v
Introduction ............................................................................................................................. vii
Amborellaceae............................................................................................................................1
Amborella ............................................................................................................................ 3
Maps...........................................................................................................................................5
Index...........................................................................................................................................7
iii

CONTRIBUTORS TO THIS PART
Text
Joël Jérémie, Muséum National d’Histoire Naturelle, Département Milieux et Peuplements
Aquatiques, USM 403, and Département Systématique et Evolution, USM 602, CP 39
(Phanérogamie), rue Cuvier 57, 75231 Paris Cedex 05, France. E-mail: jeremie@mnhn.fr
Porter P. Lowry II, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299 (USA);
and Muséum National d’Histoire Naturelle, Département Systématique et Evolution, USM 602,
CP 39 (Phanérogamie), rue Cuvier 57, 75231 Paris Cedex 05, France. E-mail: pete.lowry@mobot.org
Frédéric Tronchet, Muséum National d’Histoire Naturelle, Département Systématique et
Evolution, USM 602, CP 39 (Phanérogamie), rue Cuvier 57, 75231 Paris Cedex 05, France.
E-mail: tronchet@mnhn.fr
Illustrations
Hélène Lamourdedieu, Muséum National d’Histoire Naturelle, Département Systématique et
Evolution, USM 602, CP 39 (Phanérogamie), rue Cuvier 57, 75231 Paris Cedex 05, France
iv

Species Plantarum Steering Committee (as on December 2007)
G. Achoundong, Herbier National de Cameroon, Yaoundé, Cameroon
C.E. Anderson, University of Michigan, Ann Arbor, USA (Editor)
A. Anton, Museo Botánico, Universidad Nacional de Córdoba, Argentina
W. Berendsohn, Botanischer Garten und Botanisches Museum, Berlin, Germany
D.E. Boufford, Harvard University Herbaria, Cambridge, Mass., USA
L. Boulos, Dokki, Cairo, Egypt
R.K. Brummitt, Herbarium, Royal Botanic Gardens, Kew, UK
S. Castroviejo, Real Jardín Botánico, CSIC, Madrid, Spain
A.C. Chikuni, Royal Norwegian Embassy, Lilongwe, Malawi
M.D. Correa, Universidad de Panama, Panama
T.F. Daniel, California Academy of Sciences, San Francisco, Calif., USA
D. Darnaedi, Kebun Raya Indonesia, Bogor, Indonesia
P. Dávila, Escuela Nacional de Estudios Profesionales, UNAM, Tlalnepantla, Mexico
B. Ertter, Berkeley, California, USA
T.S. Filgueiras, Reserva Ecológica do IBGE, Brasilia, Brazil
E. Forero, Universidad Nacional de Colombia, Bogotá, Colombia
I. Friis, Botanical Museum and Library, Copenhagen, Denmark
V. Funk, Smithsonian Institution, Washington DC, USA
D. Geltman, Komarov Botanical Institute, St. Petersberg, Russia
A.S. George, Kardinya, WA, Australia
W. Greuter, Botanischer Garten und Botanisches Museum, Berlin, Germany
C. Hilton-Taylor, I.U.C.N., Cambridge, UK
Hong D.Y., Chinese Academy of Sciences, Beijing, China
K. Iwatsuki, Yokohama, Japan
B. Jonsell, Uppsala, Sweden
B. Khayota, National Museums of Kenya, Nairobi, Kenya
J. Kirschner, Inst. Bot., Pr ° uhonice, Czech Republic (Vice-Chairman, Receiving Editor)
H.P. Linder, Institut für Systematische Botanik, Zurich, Switzerland
P.-A. Loizeau, Conservatoire et Jardin botaniques, Genève, Switzerland
J.L. Luteyn, New York Botanic Gardens, Bronx, New York, USA
K. Marhold, Slovak Academy of Sciences, Bratislava, Slovakia (Editor)
M. Moraes, Universidad Mayor de San Andreas, La Paz, Bolivia
P. Morat, Muséum National d´Histoire Naturelle, Paris, France
N.R. Morin, The Arboretum at Flagstaff, Arizona, USA
Nguyen T.H., Institute of Ecology & Biological Resources, Hanoi, Vietnam
H. Ohashi, Tohoku University, Sendai, Japan
R.J. Pankhurst, Royal Botanic Garden, Edinburgh, UK
G. Prance, Royal Botanic Gardens, Kew, UK
M. Sanjappa, Botanical Survey of India, Kolkata, India
T. Santisuk, Royal Forest Department, Bangkok, Thailand
Sebsebe Demissew, Addis Ababa University, Addis Ababa, Ethiopia
J.H. Seyani, National Herbarium and Botanic Gardens, Zomba, Malawi
G.F. Smith, National Botanical Institute, Pretoria, South Africa (Chairman)
W. Thomas, New York Botanic Gardens, Bronx, New York, USA (Secretary)
W.L. Wagner, Smithsonian Institution, Washington DC, USA (Treasurer)
A.J.G. Wilson, Australian Biological Resources Study, Canberra, ACT, Australia
K. Wilson, Royal Botanic Gardens, Sydney, NSW, Australia
J.L. Zarucchi, Missouri Botanical Garden, Saint Louis, Missouri, USA
v

INTRODUCTION
Species Plantarum aims to provide in concise format, and with standardised data fields, basic
taxonomic information on the vascular plants of the world, including accepted names and
synonyms with bibliographic data, types of names, keys and descriptions from family to varietal
levels, geographical distributions, ecological information and other related matters, and to publish
it in both hard copy and electronic form.
The format of the Species Plantarum is based on that of Flora of Australia, with some departures
made necessary by the different scale of the project. Initially (vols. 1 to 11, and the introductory
volume), the series was edited and published for the Species Plantarum Project and IOPI by the
Australian Biological Resources Study (ABRS), producers of Flora of Australia. The editorial
work is being continued by J. Kirschner (receiving editor, Institute of Botany, Academy of
Sciences of the Czech Republic), C.E. Anderson (University of Michigan, Ann Arbor) and
K. Marhold (Instute of Botany, Slovak Academy of Sciences).
Treatments are contributed on a voluntary basis. Each part of Species Plantarum is intended to
provide a complete account of a family, subfamily, large genus or other related taxonomic group.
While treatments of small families may be shorter, it is intended that contributions will, in
general, cover at least 50 to 100 species. Authors should contact the receiving editor before the
preparation of their contributions.
Distribution maps are based on those in R.K.Brummitt, World Geographical Scheme for Recording
Plant Distributions 2nd edn (2001), and the ´countries´adopted are those of Level 3 and
4 that work. Description of distribution follows the same work, with a two-digit code for regions
and a three-letter code for the ´country´. Upper case letters for the ´country´indicate native
distribution; lower case letters indicate that the taxon is only present in that ´country´ as an introduced
and naturalised plant. If a taxon is extinct in a ´country´, this is indicated by a dagger.
Distribution of species as cultivated plants is not included.
Journal titles are abbreviated according to G.D.R. Bridson & E.R.Smith, Botanico-Periodicum-
Huntianum/Suplementum (1991). Book titles are abbreviated according to F.A.Stafleu &
R.S.Cowan, Taxonomic Literature (2nd edn) Vols 1-7, and Supplements (1976-), except that
upper case initial letters are used for proper names and significant words. Authors of plant names
are abbreviated according to R.K.Brummitt & C.E.Powell, Authors of Plant Names (1992).
J. Kirschner
Pru°honice
December 2007
vii

AMBORELLACEAE
J. Jérémie 1 , P.P. Lowry II 2 & F. Tronchet 3
Amborellaceae Pichon, Bull. Mus. Natl. Hist. Nat., sér. 2, 20: 384 (1948).
Type: Amborella Baill.
Trees to small trees, dioecious. Stems lacking vessels (wood homoxylous); internal phloem
absent. Leaves simple, alternate, stipules lacking. Inflorescences axillary, borne in the axils of
the leaves and on leafless branches, forming cymules, appearing fascicled, with bracts at the
base. Flowers with a pedicellate receptacle; perianth undifferentiated (tepals), acyclic; bracteoles
borne in spirals on the receptacle and sometimes grading into tepals. Staminate flowers with
stamens in spirals on the internal face of the receptacle, stamens more or less laminar, dehiscing
by longitudinal slits, introrse. Pollen grains oblate to sphaeroidal, monoaperturate (ulcerate
at distal pole), exine characterized by ektexinous tectal elements (so-called cupulae). Pistillate
flowers with free carpels borne in the center of the receptacle; stigma sessile; ovule 1, orthotropous,
pendant, micropyle inferior. Fruits: fruiting carpels drupaceous, indehiscent, each enclosing
a pendant seed, endosperm present, embryo basal.
This family is represented by a single species, Amborella trichopoda Baill., endemic to New
Caledonia.
Amborella was generally placed in the family Monimiaceae Juss. before it was recognized as
belonging to a separate family by Pichon in 1948, based on several distinctive characters:
homoxylous wood; alternate leaves; and an orthotropous, pendant ovule with an inferior
micropyle. Cladistic studies conducted at the end of the 20 th Century using molecular data resulted
in new phylogenetic trees for the angiosperms that confirmed the distinctiveness of the family
[see Soltis et al. (1997), Chase et al. (1993), APG II (2003), Qui et al. (1999), Mathews &
Donoghue (1999)]. In these works and subsequent molecular studies and discussions [e.g.,
Parkinson et al. (1999), Aoki et al. (2004), Lockhart & Penny (2005)], Amborella trichopoda
appears to represent the sister group of all other extant angiosperm taxa and can thus be considered
to represent the most ancient surviving flowering plant lineage. The putative sister position
of Amborella to all the other extant angiosperms was recently challenged by Goremykin et
al. (2003). Independently, however, Degtjareva et al. (2004) and Soltis & Soltis (2004) showed
that Goremykin’s misinterpretation is the result of inadequate taxon sampling.
The recent interest focused on Amborellaceae has resulted in a number of detailed studies
analyzing various aspects of their morphology, anatomy and development. Vegetative anatomy
was studied by Feild et al. (2000) and Carlquist & Schneider (2001), floral and inflorescence
development by Posluszny & Tomlinson (2003) and Buzgo et al. (2004), developmental
morphology of the ovules by Yamada et al. (2001), endosperm by Floyd & Friedman (2001),
embrylogy by Tobe et al. (2000), and fruit structure by Bobrov et al. (2005). The karyotype of
Amborella was investigated by Oginuma et al. (2000), and genome size by Leitch & Hanson
(2002). Reproductive structures of Amborella were analyzed by Endress & Igersheim (2000),
and floral biology and population features of the genus were elucidated by Thien et al. (2003).
A horizontal transfer of mitochondrial genes to Amborella was documented by Bergthorsson et
al. (2004).
1
Muséum National d’Histoire Naturelle, Département Milieux et Peuplements Aquatiques, USM 403, and Département
Systématique et Evolution, USM 602, CP 39 (Phanérogamie) rue Cuvier, 57, 75231 Paris Cedex 05, France.
1
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299 (USA); and Muséum National d’Histoire
Naturelle, Département Systématique et Evolution, USM 602, CP 39 (Phanérogamie) rue Cuvier, 57, 75231 Paris
Cedex 05, France.
3
Muséum National d’Histoire Naturelle, Département Systématique et Evolution, USM 602, CP 39 (Phanérogamie)
rue Cuvier 57, 75231 Paris Cedex 05, France.
1

SPECIES PLANTARUM – FLORA OF THE WORLD 2008
L.L.Money, I.W.Bailey & B.G.L.Swamy, The morphology and relationships of the Monimiaceae,
J. Arnold Arbor. 31: 372-404 (1950); J.Jérémie, Amborellaceae, in Fl. Nouvelle-
Calédonie & Dépend. 11: 157-160 (1982); F.B.Sampson, Pollen morphology of the Amborellaceae
and Hortoniaceae (Hortonioideae: Monimiaceae), Grana 32: 154-162 (1993); M.W.Chase
et al., Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene
rbcL, Ann. Missouri Bot. Gard. 80: 526-580 (1993); S.Mathews & M.J.Donoghue, The root of
angiosperm phylogeny inferred from duplicate phytochrome genes, Science 286: 947-950 (1999);
Y.-L.Qui, J.Lee, F.Bernasconi-Quadroni, D.E.Soltis, P.S.Soltis, M.Zanis, E.A.Zimmer, Z.Chen,
V.Savolainen & M.W.Chase, The earliest angiosperms: Evidence from mitochondrial, plastid,
and nuclear genomes, Nature 402: 404-407 (1999); C.L.Parkinson, K.L.Adams & J.D.Palmer,
Multigene analyses identify the three earliest lineages of extant flowering plants, Current Biol.
9: 1485-1488 (1999); P.K.Endress & A.Igersheim, The reproductive structures of the basal
angiosperm Amborella trichopoda (Amborellaceae), Int. J. Pl. Sci. 161: suppl., pp. 237-248
(2000); T.S.Feild, M.A.Zweiniecki, T.Brodribb, T.Jaffré, M.J.Donoghue & N.M.Holbrook,
Structure and function of tracheary elements in Amborella trichopoda, Int. J. Pl. Sci. 161: 705-
712 (2000); H.Tobe, T.Jaffré & P.H.Raven, Embryology of Amborella (Amborellaceae): descriptions
and polarity of character states, J. Pl. Res. 113: 271-280 (2000); K.Oginuma, T.Jaffré &
H.Tobe, The karyotype analysis of somatic chromosomes in Amborella trichopoda (Amborellaceae),
J. Pl. Res. 113: 281-283 (2000); T.Yamada, H.Tobe, R.Imaichi & M.Kato, Developmental
morphology of the ovules of Amborella trichopoda (Amborellaceae) and Chloranthus
serratus (Chloranthaceae), Bot. J. Linn. Soc. 137: 277-290 (2001); M.Hesse, Pollen characters
of Amborella trichopoda (Amborellaceae): a reinvestigation, Int. J. Pl. Sci. 162: 201-208 (2001);
S.L.Carlquist & E.L.Schneider, Vegetative anatomy of the New Caledonian endemic Amborella
trichopoda: relationships with the Illiciales and implications for vessel origin, Pac. Sci. 55: 305-
312 (2001); S.K.Floyd & W.E.Friedman, Developmental evolution of endosperm in basal
angiosperms: evidence from Amborella (Amborellaceae), Nuphar (Nymphaeaceae), and Illicium
(Illiaceae), Pl. Syst. Evol. 228: 153-169 (2001); I.J.Leitch & L.Hanson, DNA C-values
in seven families fill phylogenetic gaps in the basal angiosperms, Bot. J. Linn. Soc. 140: 175-
179 (2002); U.Posluszny & P.B.Tomlinson, Aspects of inflorescence and floral development in
the putative basal angiosperm Amborella trichopoda (Amborellaceae), Canad. J. Bot. 81: 28-
39 (2003); V.V.Goremykin, K.I.Hirsch-Ernst, S.Wolfl, F.H.Hellwig, Analysis of the Amborella
trichopoda chloroplast genome sequence suggests that Amborella is not a basal angiosperm,
Molec. Biol. Evol. 20: 1499-1505 (2003); APG II [M.W.Chase et al.], An update of the
Angiosperm Phylogeny Group classification for the orders and families of flowering plants:
APG II, Bot. J. Linn. Soc. 141: 399-436 (2003); L.B.Thien, T.L.Sage, T.Jaffré, P.Bernhardt,
V.Pontieri, P.H.Weston, D.Malloch, H.Azuma, S.W.Graham, M.A.McPherson, H.S.Rai, R.F.Sage
& J.-L.Dupré, The population structure and floral biology of Amborella trichopoda (Amborellaceae),
Ann. Missouri Bot. Gard. 90: 466-490 (2003); G.V.Degtjareva, T.H.Samigullin,
D.D.Sokoloff & C.M.Valiejo-Roman, Gene sampling versus taxon sampling: is Amborella
(Amborellaceae) a sister group to all other extant angiosperms?, Bot. Zhurn. 89: 896-907 (2004);
D.E.Soltis & P.S.Soltis, Amborella not a “basal angiosperm”? Not so fast, Amer. J. Bot. 91: 997-
1001 (2004); U.Bergthorsson, A.O.Richardson, G.J.Young, L.R.Goertzen & J.D.Palmer,
Massive horizontal transfer of mitochondrial genes from diverse land plant donors to the basal
angiosperm Amborella, Proc. Natl. Acad. Sci. U.S.A. 101: 17747-17752 (2004); S.Aoki,
K.Uehara, M.Imafuku, M.Hasebe & M.Ito, Phylogeny and divergence of basal angiosperms
inferred from APETALA3- and PISTILLATA-like MADS-box genes, J. Pl. Res. 117: 229-244
(2004); M.Buzgo, P.S.Soltis & D.E.Soltis, Floral developmental morphology of Amborella trichopoda
(Amborellaceae), Int. J. Pl. Sci. 165: 925-947 (2004); A.V.F.C.Bobrov, P.K.Endress,
A.P.Melikian, M.S.Romanov, A.N.Sorokin & A.P.Bejerano, Fruit structure of Amborella trichopoda
(Amborellaceae), Bot. J. Linn. Soc. 148: 265-274 (2005); P.J.Lockhart & D.Penny, The
place of Amborella within the radiation of angiosperms, Trends Pl. Sci. 10: 201-202 (2005).
2

AMBORELLACEAE
AMBORELLA
Amborella Baill., Hist. Pl. 1: 328 (1869).
Type: A. trichopoda Baill.
Monotypic genus, endemic to New Caledonia.
Amborella trichopoda Baill., Hist. Pl. 1: 328 (1869)
T: New Caledonia, Mt. Arago, Balansa 1800; lecto: P, fide J.Jérémie, in Fl. Nouvelle-Calédonie & Dépend. 11: 158
(1982).
Illustrations: J.Jérémie, in Fl. Nouvelle-Calédonie & Dépend. 11: 159 (1982); F.B.Sampson, Grana 32: 158 (1993);
P.K.Endress, Pl. Syst. Evol. 192: 85 (1994); P.K.Endress, Vierteljahrsschr. Naturf. Ges. Zürich 144: 10 (1999).
Map: J.Jérémie, in Fl. Nouvelle-Calédonie & Dépend. 11: 158 (1982).
Scrambling unisexual tree 2-6 m tall (species dioecious); tree habit achieved by overtopping of
older axes by basally produced younger ones. Stems cylindrical, when young with erect uni- or
multicellular trichomes. Leaves dark green, shiny above, light green below; petiole 6-10 mm
long; blade ovate to ovate-lanceolate, densely pubescent to glabrous, (6-) 8-15 cm long, (2-) 3-
8 cm wide; margins undulate to dentate (6-12 teeth per side); apex rounded to acute, often apiculate;
base rounded or truncate to subcordate; secondary nerves 6-10 pairs, generally visible
below. Inflorescence with 2-30 flowers, 1.5-6 cm long. Flowers small, unisexual; pedicel 2-12
mm long; tepals 4-8, c. 2 mm long. Staminate flowers pale yellow; stamens sessile, (6-) 11-14
(-21), inserted in a spiral on the receptacle, 1.5-1.8 mm long, triangular, the connective extending
into a small apiculum. Pistillate flowers slightly larger, white to pale yellow, carpels (3-) 5-
8, free, c. 2 mm long (often 1-2 tepals bearing a sterile anther); ovule 0.5-0.7 mm long. Fruits
red at maturity, stipitate, ± ellipsoid, slightly flattened, 8-10 mm long, 4-5 mm wide. Seed c. 8
mm long; endocarp lignified, evidently rugose. 2n = 26, fide K.Oginuma, T.Jaffré & H.Tobe,
J. Pl. Res. 113: 281-283 (2000), genome size small, 1C = 0.89 pg, fide I.J.Leitch & L.Hanson,
Bot. J. Linn. Soc. 140: 175-179 (2002). Fig. 1 and front cover drawing.
Locally abundant in central New Caledonia between 200 and 1,000 m altitude. 60: NWC. Dense,
humid forest and gallery forest, often in slightly disturbed areas mainly on schist substrates.
Staminate plants prevail in natural populations, flowering is synchronised, wind and insect
pollination has been observed. Amborella is a generalist as regards insect pollinators, for which
pollen represents a reward (nectar has not been observed). Maps. 1 & 2.
60. NEW CALEDONIA: Mt. Arago, 700 m, B.Balansa 1800 (P); Koindé, 800-900 m, L.Bernardi 9959 (G, P);
Mt. Dogny, 800 m, J Jérémie & C.Tirel 1587 (MO, NOU, P); Plateau de Dogny, 870 m, P.P.Lowry 3386 (MO, NOU, P);
Tiwaka, Moindip (pente est), 700 m, H.S.MacKee 28460 (MO, NOU, P); NNE Sarraméa, 750 m, G.McPherson 2607
(MO, NOU, P); Mt. Aoupinié, 850 m, Ph.Morat 5083 (NOU, P); Nakada, 600 m, Ph.Morat 6533 (NOU, P);
Mt. Pembaï, N col d’Amieu, SW route Canala-La Foa, 830 m, R.B.Phillips & M.Schmid 3193 (NOU, P); Table Unio,
650 m, R.Schodde 5245 (A, AD, CANB, L, P).
3

SPECIES PLANTARUM – FLORA OF THE WORLD 2008
Figure 1. Amborella trichopoda Baill. A, flowering branch; B, floral bud; C, stamen prior
to anthesis; D, staminate flower; E, dehisced stamen; F, pistillate flower; G, carpel in cross
section; H, inflorescences branch; J, fruit; K, seed. Scale bars: A, H = 1 cm; B, J = 2 mm;
C, E, G = 0.5 mm; D, F, K = 1 mm. Reproduced with permission, from Fl. Nouvelle-Calédonie
& Dépend. 11: 159 (1982). Drawn by H. Lamourdedieu.
4

Maps

SPECIES PLANTARUM – FLORA OF THE WORLD 2008
1. Amborella trichopoda (world distribution)
2. Amborella trichopoda (distribution in New Caledonia)
6

INDEX
Synonyms are in italics. Page numbers in boldface indicate primary page references. Page
numbers with an asterisk (*) indicate pages with illustrations or maps.
Amborellaceae Pichon 1, 2
Amborella Baill. 1, 2, 3
trichopoda Baill. 1, 2, 3, 4*, 6*
Monimiaceae Juss. 1
7

Parts already published in this series:
Introduction to the series (English, 1999)
Introduction to the series (Chinese, 2000)
1. Irvingiaceae, 1999
2. Stangeriaceae, 1999
3. Welwitschiaceae, 1999
4. Schisandraceae, 2001
5. Prioniaceae, 2001
6. Juncaceae 1: Rostkovia to Luzula, 2002
7. Juncaceae 2: Juncus subg. Juncus, 2002
8. Juncaceae 3: Juncus subg. Agathryon, 2002
9. Chrysobalanaceae 1: Chrysobalanus to Parinari, 2003
10. Chrysobalanaceae 2: Acioa to Magnistipula, 2003
11. Saururaceae, 2005
12. Opiliaceae, 2008
13. Paracryphiaceae, 2008