Table Of ContentTHE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K.Kubitzki
Springer-Verlag Berlin HeidelbergGmbH
Volumes published in this series:
Volume I Pteridophytes and Gymnosperms
Edited by K.U.Kramer and P.S.Green (1990)
Volume II Flowering Plants.Dicotyledons.Magnoliid,Hamamelid and Caryophyllid Families
Edited by K.Kubitzki,J.G.Rohwer,and V.Bittrich (1993)
Volume IIIFlowering Plants.Monocotyledons:Lilianae (except Orchidaceae)
Edited by K.Kubitzki (1998)
Volume IVFlowering Plants.Monocotyledons:Alismatanae and Commelinanae
(except Gramineae)
Edited by K.Kubitzki (1998)
Volume V Flowering Plants.Dicotyledons:Malvales,Capparales and Non-betalain
Caryophyllales
Edited by K.Kubitzki and C.Bayer (2003)
The Families
and Genera
of Vascular Plants
Edited by K.Kubitzki
V
Flowering Plants · Dicotyledons
Malvales, Capparales and Non-betalain Caryophyllales
Volume Editors:
K. Kubitzki and C. Bayer
With 97 Figures
1 3
Professor Dr.KlausKubitzki
Institut für Allgemeine Botanik
Ohnhorststraße 18
22609 Hamburg
Germany
Dr.ClemensBayer
Palmengarten
Siesmayerstraße 61
60323 Frankfurt am Main
Germany
Library ofCongress-in-Publication Data
Flowering plants,Dicotyledons :Malvales,Capparales,and non-betalain Caryophyllales/
volume editors,K.Kubitzki and C.Bayer.
p. cm.– (The Families and genera ofvascular plants ;5)
Includes bibliographical references.
ISBN978-3-642-07680-0 ISBN978-3-662-07255-4(eBook)
DOI 10.1007/978-3-662-07255-4
1. Dicotyledons – Classification. I Kubitzki,Klaus,1933– II. Bayer,Clemens. III. Series.
OK495.A12F62 2002
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Preface
When more than two decades ago this series was planned,classification at the level of
higher categories was rather contentious and considered to be of little concern for this
book series.Through the advancement of molecular systematics within the last decade,
the outlines ofa phylogenetically based family scheme ofangiosperms have become avail-
able.This phylogenetic scheme for the first time made meaningful broader comparisons
feasible and is adopted here as a guideline for the circumscription of major blocks of
families.However,due to the fragility of formal higher level classification only one cate-
gory above the rank of family,that of order,will be used in this volume.
Despite the great progress in the understanding of broader phylogenetic relationships
made during the last years,the circumscription of many families,let alone of genera,is
still far from being settled and,in the contributions to this volume,the arrangement of
genera within families is often phenetic rather than phylogenetic.This difficulty is par-
ticularly acute in larger families,and despite much molecular work that has been con-
ducted in Cruciferae,the authors of that family account to this volume felt it premature
to provide a formal subdivision of the group.Also,in other families,and particularly in
Malvaceae-Malvoideae, the arrangement and circumscription of genera, for lack of a
better alternative,are largely phenetic.Nevertheless,it is hoped that the classification pre-
sented here is an advance over what had been known previously.
One family which remains unclassified as to order, Tapisciaceae, is included in this
volume, because in various molecular analyses it appeared sister to Capparales. For
another family,Emblingiaceae,equivocal results had been obtained by molecular analy-
ses but its place in Capparales,originally suggested by F.von Mueller,has now been con-
firmed. This enabled us to include the family into this volume at a late date of its
production but also explains why Emblingiaceae are not mentioned in the introduction
to and synopsis of capparalean families.
Apart from the unselfish collaboration by the authors,the preparation of this volume
has received invaluable support by several individuals,to whom I wish to extend my deep
gratitude.Victor Badillo,David Bates,David Baum,Richard Brummitt,Sherwin Carlquist,
Mark J.E.Coode,Paul Fryxell,Owi Nandi,Mark E.Olson,J.E.Rodman,George E.Schatz,
and Rüdiger Seine read manuscripts or provided access to unpublished information.Mark
W.Chase,Vincent Savolainen,and Peter F.Stevens shared with me their intimate knowl-
edge ofrecent developments in angiosperm systematics.Peter H.Raven is thanked for his
efforts in making a treatment ofCruciferae for this series possible.Special thanks are due
to Susanne S.Renner,who over many years has acted as an intermediary to many,mostly
North American,authors.
Finally,I would like to express my gratitude to all copyright holders for their generous
permission to take over illustrations under their copyright in the present volume.Col-
laboration with the staff of Springer Verlag was pleasant,as always.
Hamburg,June 2002 K.Kubitzki
Contents
Introduction to Expanded Caryophyllales
Ph.Cuénoud . . . . . . . . . . . . . . . . . . . . . . . 1
Conspectus ofFamilies ofExpanded Caryophyllales
Ph.Cuénoud . . . . . . . . . . . . . . . . . . . . . . . 5
Introduction to Capparales
K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 7
Conspectus ofthe Families ofCapparales
K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 11
Introduction to Malvales
K.Kubitzkiand M.W.Chase . . . . . . . . . . . . . . 12
Conspectus ofthe Families ofMalvales
K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 17
GENERAL REFERENCES . . . . . . . . . . . . . . . . . 19
Akaniaceae C.Bayerand O.Appel . . . . . . . . . . . . . . . . . . 21
Ancistrocladaceae S.Porembski . . . . . . . . . . . . . . . . . . . . . . . 25
Asteropeiaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 28
Bataceae C.Bayerand O.Appel . . . . . . . . . . . . . . . . . . 30
Bixaceae H.-H.Poppendieck . . . . . . . . . . . . . . . . . . . . 33
Capparaceae L.E.Kers . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Caricaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 57
Cistaceae J.M.Arringtonand K.Kubitzki . . . . . . . . . . . . 62
Cochlospermaceae H.-H.Poppendieck . . . . . . . . . . . . . . . . . . . . 71
Cruciferae O.Appeland I.Al-Shehbaz . . . . . . . . . . . . . . . 75
Diegodendraceae C.Bayer . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Dioncophyllaceae S.Porembskiand W.Barthlott . . . . . . . . . . . . 178
Dipterocarpaceae P.S.Ashton . . . . . . . . . . . . . . . . . . . . . . . . 182
Droseraceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 198
Drosophyllaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 203
Emblingiaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 206
Frankeniaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 209
Gyrostemonaceae A.S.George . . . . . . . . . . . . . . . . . . . . . . . . 213
VIII Contents
Koeberliniaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 218
Limnanthaceae C.Bayerand O.Appel . . . . . . . . . . . . . . . . . . 220
Malvaceae C.Bayerand K.Kubitzki . . . . . . . . . . . . . . . . 225
Moringaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 312
Muntingiaceae C.Bayer . . . . . . . . . . . . . . . . . . . . . . . . . . 315
Nepenthaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 320
Neuradaceae C.Bayer . . . . . . . . . . . . . . . . . . . . . . . . . . 325
Pentadiplandraceae C.Bayerand O.Appel . . . . . . . . . . . . . . . . . . 329
Physenaceae W.C.Dickison† . . . . . . . . . . . . . . . . . . . . . . 332
Resedaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 334
Rhabdodendraceae G.T.Prance . . . . . . . . . . . . . . . . . . . . . . . . 339
Salvadoraceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 342
Sarcolaenaceae C.Bayer . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Setchellanthaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 353
Simmondsiaceae E.Köhler . . . . . . . . . . . . . . . . . . . . . . . . . 355
Sphaerosepalaceae C.Bayer . . . . . . . . . . . . . . . . . . . . . . . . . . 359
Tamaricaceae J.F.Gaskin . . . . . . . . . . . . . . . . . . . . . . . . . 363
Tapisciaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 369
Tepuianthaceae K.Kubitzki . . . . . . . . . . . . . . . . . . . . . . . . 371
Thymelaeaceae B.E.Herber . . . . . . . . . . . . . . . . . . . . . . . . 373
Tovariaceae O.Appeland C.Bayer . . . . . . . . . . . . . . . . . . 397
Tropaeolaceae C.Bayerand O.Appel . . . . . . . . . . . . . . . . . . 400
Addendumto Volume II
Hydrastidaceae H.Tobe . . . . . . . . . . . . . . . . . . . . . . . . . . 405
Index ofScientific Names . . . . . . . . . . . . . . . . . 411
List of Contributors
Al-Shehbaz,I.A. Missouri Botanical Garden,P.O.Box 299,St.Louis,
Missouri 64507,USA
Appel,O. Bredkamp 36e,22589 Hamburg,Germany
Arrington,J.M. Department of Biology,212 Mueller Laboratory,Pennsylvania
State University,University Park,Pennsylvania 16802,USA
Ashton,P.S. Arnold Arboretum of Harvard University,22 Divinity
Avenue,Cambridge,Massachusetts 02138,USA
Barthlott,W. Botanisches Institut,Universität Bonn,
Meckenheimer Allee 170,53115 Bonn,Germany
Bayer,C. Palmengarten,Siesmayerstr.61,60323 Frankfurt am Main,
Germany
Chase,M.W. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AS,UK
Cuénoud,Ph. 65 Bois de la Chapelle,1213 Onex,Switzerland
Dickison,W.C. deceased
Gaskin,J.F. Missouri Botanical Garden,P.O.Box 299,St.Louis,
Missouri 64507,USA
George,A.S. “Four Gables”,18 Barclay Rd.,Kardiny,W.A.6163,Australia
Herber,B.E. Institut für Allgemeine Botanik,Universität Hamburg,
22609 Hamburg,Germany
Kers,L.E. Bergius Botanic Garden,P.O.B.50017,10405 Stockholm,
Sweden
Koehler,E. Spezielle Botanik und Arboretum,Humboldt-Universität,
Späthstr.80/81,12437 Berlin,Germany
Kubitzki,K. Institut für Allgemeine Botanik,Universität Hamburg,
22609 Hamburg,Germany
Poppendieck,H.-H. Institut für Allgemeine Botanik,Universität Hamburg,
22609 Hamburg,Germany
Porembski,S. Institut für Allgemeine und Spezielle Botanik,
Universität Rostock,Wismarische Str.8,18057 Rostock,
Germany
Prance,G.T. The Old Vicarage,Silver Street,Lyme Regis,
Dorset DT7 3HS,UK
Tobe,H. Department of Botany,Graduate School of Science,
Kyoto University,Kyoto 606-8502,Japan
Introduction to Expanded Caryophyllales 1
Introduction to Expanded Caryophyllales1
Ph.Cuénoud
The order Caryophyllales (s.str., also called Cen- and forming, together with Caryophyllales, the
trospermae) has been identified as a natural group subclass Caryophyllidae.
by early workers (Braun 1864; Eichler 1876).The Molecular systematic studies (starting,i.a.with
main characters used for the recognition of this the work ofRettig et al.1992 and Chase et al.1993)
group include the free-central or basal placenta- have provided an additional,independent source
tion, the presence of perisperm instead of of information in favour of the delimitation of
endosperm as seed storage tissue, and curved Caryophyllales. Subsequent molecular studies,
embryos (Bittrich 1993).More recent studies have such as those by Morton et al.(1997) and Fay et al.
provided further support for the naturalness of (1997), have added numerous further families to
this group.Thus,all Caryophyllales families (with this group, which now is thought to include, in
the exception of Caryophyllaceae and Mollugi- addition to Polygonaceae and Plumbaginaceae,
naceae) synthesise the unique betalain pigments the carnivorous families Dioncophyllaceae,
(for review see Clement et al. 1994) instead of Droseraceae, Drosophyllaceae, and Nepenthaceae
anthocyanins,which are ubiquitous in all remain- with the closely related Ancistrocladaceae, two
ing seed plants and ferns. Caryophyllales also Malagasy families, Asteropeiaceae and Physe-
possess bound ferulic acid in unlignified cell- naceae, the neotropical Rhabdodendraceae, the
walls, a characteristic otherwise known only of North American monospecific Simmondsiaceae,
some monocotyledons (Hartley and Harris 1981). and the widespread Frankeniaceae and Tamari-
Ultrastructural studies have revealed that caceae (Fig. 1). The broadened understanding of
Caryophyllales share a unique type of sieve- the phylogenetic position ofthese families led the
element plastids (P3; Behnke 1994), in which a Angiosperm Phylogeny Group (APG 1998) to
peripheral ring of proteinaceous filaments sur- introduce the concept of “expanded Caryophyl-
rounds a protein crystal (sometimes absent) of lales”,which now comprises 26 families,the tradi-
either globular or angular shape.Based on the data tionally recognised Centrospermae often being
available by then, Cronquist and Thorne (1994) referred to as “core Caryophyllales”. Significant
circumscribed the order Caryophyllales to characters of the “expanded Caryophyllales”
comprise 11 families: Aizoaceae, Amarantha- include anomalous secondary growth, spinulose
ceae, Basellaceae, Cactaceae, Caryophyllaceae, pollen sexine,at most biseriate wood rays,persist-
Chenopodiaceae (including Sarcobataceae and ent calyces, alternate vessel pitting, a starchy
Halophytaceae), Didiereaceae, Molluginaceae, endosperm (not applying to core Caryophyllales),
Nyctaginaceae, Phytolaccaceae (including Agdes- lack of a hypostase,and lack of mucilage cavities
tidaceae, Achatocarpaceae, Barbeuiaceae, Giseki- (Nandi et al.1998; Savolainen,Chase et al.2000).
aceae, Petiveriaceae, and Stegnospermataceae), Many members of Caryophyllales show adapta-
and Portulacaceae (including Hectorellaceae). In tions to extreme habitats. Aizoaceae, Cactaceae,
his classification, Cronquist (1981) considered Didiereaceae, some Chenopodiaceae and others
Polygonaceae and Plumbaginaceae as the closest withstand xeric conditions. Frankeniaceae, Ta-
relatives of Caryophyllales,each in its own order maricaceae,Plumbaginaceae and some Chenopo-
diaceae resist salinity, and the carnivorous
Droseraceae, Drosophyllaceae, and Nepenthaceae
are adapted to nitrogen-poor soils.
1 The “classical”(betalain-producing) Caryophyllales or “core Within the expanded Caryophyllales,according
Caryophyllales”had been treated in 1993 in Volume II ofthis to molecular data (e.g.Savolainen,Fay et al.2000),
series (cf.Kubitzki et al.1993).At that time,very few results of core Caryophyllales form a clade closely related to
molecular systematic studies were available and here the
Asteropeiaceae and Physenaceae.With the excep-
opportunity is taken to review their phylogenetic relation-
tion of Simmondsiaceae and Rhabdodendraceae,
ships,together with those ofthe non-betalain Caryophyllales
which since have been added to this group. whose position is not yet settled (Simmondsiaceae
2 Ph.Cuénoud
Fig. 1. Composite cladogram of extended Caryophyllales used here as outgroups). Bootstrap support *50–64%,
families,based on different taxon samplings of atpB,matK, **65–84%,***85–100%.(Data from Cuénoud et al.2002)
rbcL and 18S rDNA sequences (Dilleniaceae and Santalales are
