Flowering Plants · Dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae)

486 Pages·2004·10.3 MB·English

by  J. W. Kadereit (auth.)

THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
Springer Verlag Berlin Heidelberg GmbH
Volumes published in this series:
Volume I Pteridophytes and Gymnosperms
Edited by K.U. Krame r and P.S. Green (1990)
Date of publication: 28.9.1990
Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families
Edited by K. Kubitzki, J.G . Rohwer, and V. Bittrich (1993)
Date ofpublication: 28.7.1993
Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)
Edited by K. Kubitzki (1998)
Date of publication: 27.8.1998
Volume IV Flowering Plants. Monocotyledons: Alismatana e and Commelinanae (except
Gramineae)
Edited by K. Kubitzki (1998)
Date ofpublication: 27.8.1998
Volume V Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain
Caryophyllales
Edited by K. Kubitzki and C. Bayer (2003)
Date of publication: 12.9.2002
Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales , Rosales, Cornales, Ericales
Edited by K. Kubitzki (2004)
Date ofpublication: 21.1.2004
Volume VII Flowering Plants. Dicotyledons: Lamiale s (except Acanthaceae including
Avicenniaceae)
Edited by J.W. Kadereit (2004)
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzk i
T 7 " T T Flowering Plants • Dicotyledons
V .X. .X. Lamiales (exceptAcanthaceae includingAvicenniaceae)
Volume Editor:
J.W. Kadereit
With 60 Figures, 8 in Color
fKSpringer
TV84*!?-*
Professor Dr. KLAUS KUBITZKI
Universität Hamburg
Biozentrum Klein-Flottbek un d Botanischer Garten
Ohnhorststraße 18
22609 Hamburg
Germany
Professor Dr. JOACHIM W. KADEREIT
Johannes Gutenberg-Universität Mainz
Institut für Spezielle Botanik und Botanischer Garten
55099 Mainz
Germany
Library of Congress Cataloging-in-Publication Data
Flowering plants, dicotyledons : Lamiales (except Acanthaceae including Avicenniaceae) / volume editor, J.W. Kadereit.
p. cm. - (The families and genera of vascular plants ; 7)
Includes bibliographical references.
ISBN 978-3-642-62200-7 ISBN 978-3-642-18617-2 (eBook)
DOI 10.1007/978-3-642-18617-2
1. Lamiales - Classification. I. Kadereit, J.W. (Joachim W.), 1956- II. Series.
QK495.A12F57 2004
583'.96 - dc22 2003065479
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Preface
This volume,the seventh in this series,deals with the families ofthe Lamiales,a group of
the asterids or Sympetalae,which was recognised by early workers such as Bartling and
Lindley under the name Labiatiflorae or Personatae,and which has only moderately been
modified in circumscription over time.Nevertheless,our considerably increased knowl-
edge of the group and the results of molecular studies provide a more detailed basis for
understanding relationships among and evolution within the families of the Lamiales,
although,as J.W.Kadereit emphasises in his introduction,in this connection and also in
relation to family delimitation,many questions remain currently unanswered.
My sincere thanks go to all authors for their scholarly treatment of families in this
volume. In particular, the authors/organisers who handled the four large families with
more than 100 genera, Bignoniaceae, Gesneriaceae, Labiatae and Scrophulariaceae,
deserve special mention for their herculean work. The authors also emphasised the
various themes ofdiversification,differing from group to group,that are so important for
the understanding and interpretation of taxonomic diversity, such as phytochemical
diversification in Labiatae and pollinator interactions in Gesneriaceae,to cite only two
salient examples. In the circumscription of Scrophulariaceae, a pragmatic position has
been taken; since this series aims primarily at providing baseline information for fami-
lies and genera,the problem of a phylogenetically satisfactory classification of the Scro-
phulariaceae complex that avoids paraphyly at our present state of knowledge must be
left undecided,and indeed is not the primary focus of this volume.Sincere thanks are
also extended to Professor J.W.Kadereit for patiently bringing his work on the present
volume to an end,in spite of all discouraging vicissitudes.
I am most grateful to all copyright holders for so generously authorising the use of
their precious illustrative material.Finally,it is a pleasure to thank,also in the name of
Professor Kadereit,the staff of Springer-Verlag and ProEdit,Heidelberg,for their dedi-
cated work on the present volume.I would like to express special thanks to the copy editor,
Monique Delafontaine,for the many improvements she made to the manuscript.
Hamburg,December 2003 K.Kubitzki
Contents
Lamiales:Introduction and Conspectus
J.W.Kadereit . . . . . . . . . . . . . . . . . . . . . . . 1
Bignoniaceae E.Fischer,I.Theisenand L.G.Lohmann . . . . . . . 9
Buddlejaceae B.Oxelman,P.Kornhalland E.M.Norman . . . . . 39
Byblidaceae J.G.Conranand R.Carolin . . . . . . . . . . . . . . . 45
Callitrichaceae C.Erbarand P.Leins . . . . . . . . . . . . . . . . . . 50
Carlemanniaceae M.Thiv . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Cyclocheilaceae SebsebeDemissew . . . . . . . . . . . . . . . . . . . . 60
Gesneriaceae A.Weber . . . . . . . . . . . . . . . . . . . . . . . . . 63
Globulariaceae G.Wagenitz . . . . . . . . . . . . . . . . . . . . . . . 159
Hippuridaceae P.Leinsand C.Erbar . . . . . . . . . . . . . . . . . . 163
Labiatae R.M.Harley,S.Atkins,A.L.Budantsev,P.D.Cantino,
B.J.Conn,R.Grayer,M.M.Harley,R.de Kok,
T.Krestovskaja,R.Morales,A.J.Paton,
O.Rydingand T.Upson . . . . . . . . . . . . . . . . 167
Lentibulariaceae E.Fischer,W.Barthlott,R.Seine
and I.Theisen . . . . . . . . . . . . . . . . . . . . . 276
Martyniaceae H.-D.Ihlenfeldt . . . . . . . . . . . . . . . . . . . . . 283
Myoporaceae I.Theisenand E.Fischer . . . . . . . . . . . . . . . . 289
Nesogenaceae R.M.Harley . . . . . . . . . . . . . . . . . . . . . . . 293
Oleaceae P.S.Green . . . . . . . . . . . . . . . . . . . . . . . . . 296
Pedaliaceae H.-D.Ihlenfeldt . . . . . . . . . . . . . . . . . . . . . 307
Phrymaceae P.D.Cantino . . . . . . . . . . . . . . . . . . . . . . . 323
Plantaginaceae A.E.Schwarzbach . . . . . . . . . . . . . . . . . . . . 327
Plocospermataceae L.Struweand S.R.Jensen . . . . . . . . . . . . . . . . 330
Scrophulariaceae E.Fischer . . . . . . . . . . . . . . . . . . . . . . . . . 333
Stilbaceae H.P.Linder . . . . . . . . . . . . . . . . . . . . . . . . 433
Tetrachondraceae S.J.Wagstaff . . . . . . . . . . . . . . . . . . . . . . . 441
Trapellaceae H.-D.Ihlenfeldt . . . . . . . . . . . . . . . . . . . . . 445
Verbenaceae S.Atkins . . . . . . . . . . . . . . . . . . . . . . . . . . 449
Index to Scientific Names . . . . . . . . . . . . . . . . . 469
List of Contributors
Atkins,S. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Barthlott,W. Botanisches Institut und Botanischer Garten der Rheinischen
Friedrich-Wilhelms-Universität,Meckenheimer Allee 170,
53115 Bonn,Germany
Budantsev,A.L. Komarov Botanical Institute,Herbarium,Prof.Popov Street,
St.Petersburg 197376,Russia
Cantino,P.D. Department of Environmental and Plant Biology,Ohio
University,Athens,OH 45701-2979,USA
Carolin,R. Pulman’s Cottage,30 Pulman Street,Berry,N.S.W.2535,
Australia
Conn,B.J. Royal Botanic Garden,Sydney,Australia
Conran,J.G. Environmental Biology,School of Earth and Environmental
Sciences,The University ofAdelaide,SA 5005,Australia
Erbar,C. Heidelberger Institut für Pflanzenwissenschaften (HIP) –
Biodiversität und Pflanzensystematik,
Im Neuenheimer Feld 345,69120 Heidelberg,Germany
Fischer,E. Institut für Biologie,Universität Koblenz/Landau,
Universitätsstr.1,56070 Koblenz,Germany
Grayer,R. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Green,P.S. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Ihlenfeldt,H.-D. Ostseestr.40,24369 Waabs,Germany
Harley,M.M. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Harley,R.M. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Jensen,S.R. Department of Chemistry,The Technical University of
Denmark,2800 Lyngby,Denmark
Kadereit,J.W. Institut für Spezielle Botanik und Botanischer Garten,
Johannes Gutenberg-Universität Mainz,55099 Mainz,
Germany
Kok,R.de Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Kornhall,P. Department of Systematic Botany,Evolutionary Biology
Centre Uppsala University,Norbyv.18D,752 36 Uppsala,
Sweden
Krestovskja,T. Komarov Botanical Institute,Herbarium,Prof.Popov Street,
St.Petersburg 197376,Russia
Kubitzki,K. Institut für Allgemeine Botanik,Universität Hamburg,
Ohnhorststr.18,22609 Hamburg,Germany
List ofContributors IX
Leins,P. Heidelberger Institut für Pflanzenwissenschaften (HIP) –
Biodiversität und Pflanzensystematik,
Im Neuenheimer Feld 345,69120 Heidelberg,Germany
Linder,H.P. Institute for Systematic Botany,University of Zürich,
Zollikerstrasse 107,8008 Zürich,Switzerland
Lohmann,L.G. Missouri Botanical Garden,P.O.Box 299,St.Louis,
Missouri 64507,USA
Morales,R. Herbario,Real Jardin Botanico,Plaza de Murillo 2,
Madrid 28014,Spain
Norman,E.M. Department of Biology,Stetson University,Deland,
FL 32723,USA
Oxelman,B. Department of Systematic Botany,Evolutionary Biology
Centre Uppsala University,Norbyv.18D,752 36 Uppsala,
Sweden
Paton,A.J. Royal Botanic Gardens Kew,Richmond,Surrey TW9 3AB,UK
Ryding,O. Botanical Museum,University of Copenhagen,
Gothersgade 130,1123 Copenhagen K,Denmark
Schwarzbach,A.E. Department of Biological Sciences,Kent State University,
Kent,OH 44242,USA
Sebsebe Demissew The National Herbarium,Science Faculty,Addis Ababa
University,P.O.Box 3434,Addis Ababa,Ethiopia
Seine,R. European Astronaut Centre,Linder Höhe,
51147 Köln,Germany
Struwe,L. Department of Ecology,Evolution,and Natural Resources,
Rutgers University – Cook College,237 Foran Hall,59 Dudley
Road,New Brunswick,NJ 08901-8551,USA
Theisen,I. Botanisches Institut und Botanischer Garten der Rheinischen
Friedrich-Wilhelms-Universität,Meckenheimer Allee 170,
53115 Bonn,Germany
Thiv,M. Staatl.Museum für Naturkunde,Rosenstein 1,
70191 Stuttgart,Germany
Upson,T. Cambridge University Botanic Garden,Cory Lodge,Bateman
Street,Cambridge CB2 1JF,UK
Wagenitz,G. A.v.Haller-Institut,Abteilung Systematische Botanik,
Untere Karspüle 2,37073 Göttingen,Germany
Wagstaff,S.J. Allan Herbarium,Landcare Research,P.O.Box 69,
Lincoln 8152,New Zealand
Weber,A. Institut für Botanik,Universität Wien,Rennweg 14,
1030 Wien,Austria
Lamiales:Introduction and Conspectus 1
Lamiales: Introduction and Conspectus
J.W.Kadereit
Lamiales as presented here are a well-supported son to the latter three systems,the Lamiales as cir-
clade of the Lamiids (Bremer et al.2002) found in cumscribed here differ by the exclusion of Borag-
all major,recent molecular studies of the Asterids inaceae (incl. Lennoaceae), treated as part of
(APG II 2003; see also Wagenitz 1992). The Lamiales by Cronquist (1981),and by the inclusion
Lamiids,comprising approximately 23,600 species of Byblidaceae (Cronquist 1981: Rosales; Thorne
in 1140 genera,are herbaceous or less commonly 1992: Pittosporales; Takhtajan 1997: Aralianae-
woody plants with often opposite leaves. The Byblidales), Carlemanniaceae (Cronquist 1981:
flowers are sympetalous and mostly 5-merous, Dipsacales;Thorne 1992:not recognized;Takhta-
often representing zygomorphic lip flowers with a jan 1997:Rubiales) and Plocospermataceae (Cron-
showy corolla and with a single whorl of stamens quist 1981:Gentianales;Thorne 1992:Gentianales;
(the five stamens often reduced to four or two), Takhtajan 1997:Gentianales),treated as belonging
and a bicarpellate ovary with the two carpels in elsewhere by Cronquist (1981),Thorne (1992) and
median position (Endress 1994). Anthers very Takhtajan (1997). There is good molecular evi-
often have an outgrowth of the connective into dence for the inclusion of these three families in
the anther locule,known as pollen sac placentoid Lamiales (Byblidaceae:Hedrén et al.1995;Bremer
(Hartl 1964),glandular hairs with a head with only et al. 2001; Albach et al. 2001; Carlemanniaceae:
vertical cell walls are often present, and pollen Savolainen, Fay et al. 2000; Bremer et al. 2001;
tubes appear to lack callose (Prósperi and Cocucci Plocospermataceae: Bremer et al. 2001), and
1979;Cocucci 1983).Both micropylar and chalazal other characters in support of this can be found.
endosperm haustoria are common (Yamazaki Thus, Plocospermataceae contain cornoside as a
1974), endosperm formation is cellular, and caffeic acid derivative commonly found in the
embryogeny normally is of the Onagrad type order (Jensen 1996,2000),the carnivorous Bybli-
(Polygonad in Oleaceae; Albach et al. 2001). daceae share embryological characters with the
Protein bodies in the nuclei are widespread (Speta equally carnivorous Lentibulariaceae (Lang 1901;
1977, 1979; Bigazzi 1984, 1989a, b). Chemically, Conran 1996),and Carlemanniaceae have flowers
typical constituents of the order are 6- and/or 8- with only two stamens which may prove to be
hydroxylated flavone glycosides (Grayer et al. morphological support of their sister-group
1999), verbascosides and cornoside as charac- relationship with Oleaceae.
teristic caffeic acid derivatives (Mølgaard and The exact relationships of Lamiales within
Ravn 1988; Jensen 1992), shikimate-derived Lamiids are not finally resolved,but it is clear from
anthraquinones (Jensen 1992), C11-decarboxy- molecular evidence (e.g.Downie and Palmer 1992;
lated iridoids (Jensen 1991,1992),and stachyose or Olmstead et al.1992,1993; Backlund and Bremer
other oligosaccharides as carbon storage com- 1997;Olmstead et al.2000;Savolainen,Chase et al.
pounds (Hegnauer 1989). As will be discussed 2000;Savolainen,Fay et al.2000;Soltis et al.2000;
below, several of the characters listed above, Albach et al. 2001; Bremer et al. 2001; Olmstead
including the zygomorphic lip flowers,originated et al. 2001) that they are part of a larger clade
only within the order (Stevens 2001). containing Boraginaceae/Boraginales, Gen-
In large part, Lamiales as circumscribed here tianales, Solanales, Vahliaceae. In some of these
correspond to the Tubiflorae (Wettstein 1935; analyses (Olmstead et al.2000; Savolainen,Chase
Melchior 1964) or Personatae (Drude 1885– et al.2000) they are sister to Gentianales.Inclusion
1887; Warming and Möbius 1929; Huber 1991) ofVahliaceae into Lamiales should be considered
of earlier authors, and to Cronquist’s (1981) (Savolainen,Fay et al.2000;Albach et al.2001).
Lamiales, Callitrichales, Plantaginales and Scro- Although the Lamiales clearly represent a
phulariales,to Thorne’s (1992) Bignoniales,and to monophylum, their division into families in this
Takhtajan’s (1997) Oleales,Scrophulariales,Lami- treatment differs from that proposed by APG II
ales,Callitrichales and Hippuridales.In compari- (2003;see Table 1).
K. Kubitzki (ed.), The Families and Genera of Vascular Plants
© Springer-Verlag Berlin Heidelberg 2004
2 J.W.Kadereit
Table 1. Families recognized in this volume compared with aceae. Buddlejaceae, Callitrichaceae, Globulari-
the APG classification aceae,Hippuridaceae,Plantaginaceae s.str.as well
as Myoporaceae are treated as separate families.
APG II FGVP
Also,Phrymaceae here are treated as a monotypic
Acanthaceae Acanthaceae family. Such treatment of Scrophulariaceae s.l. is
Bignoniaceae Bignoniaceae non-phylogenetic in at least one respect.It is quite
=Scrophulariaceae s.str. Buddlejaceae clear that Scrophulariaceae s.l. are paraphyletic
Byblidaceae Byblidaceae
when Buddlejaceae, Callitrichaceae, Globulari-
=Plantaginaceae s.l. Callitrichaceae
aceae,Hippuridaceae,Myoporaceae and Plantagi-
Calceolariaceae Srophulariaceae s.l.
Carlemanniaceae Carlemanniaceae naceae,and probably also Phrymaceae are treated
Orobanchaceae Cyclocheilaceae as separate families. Buddlejaceae and Myopo-
Gesneriaceae Gesneriaceae raceae group with one part ofScrophulariaceae s.l.
==PPllaannttaaggiinnaacceeaaee ss..ll.. GHliopbpuulrairdiaacceeaaee with good support (=Scrophulariaceae s.str.:78%;
Labiatae Labiatae Olmstead et al.2001),and Callitrichaceae,Globu-
Lentibulariaceae Lentibulariaceae lariaceae and Hippuridaceae group with another
Martyniaceae Martyniaceae part of Scrophulariaceae s.l., although with only
=Scrophulariaceae s.str. Myoporaceae low support (= Plantaginaceae = Veronicaceae:
Orobanchaceae Nesogenaceae
48%; Olmstead et al. 2001). The recognition of a
Oleaceae Oleaceae
Orobanchaceae =Scrophulariaceae s.l. close relationship between most of the preceding
Paulowniaceae =Scrophulariaceae s.l. families and Scrophulariaceae is by no means
Pedaliaceae Pedaliaceae novel and had already been recognized (and partly
Phrymaceae Phrymaceae
formalized) by, for example, Hallier (1903) and
Plantaginaceae s.l. Plantaginaceae s.str.
Hartl (1965–1974). Following Olmstead et al.
Plocospermataceae Plocospermataceae
Schlegeliaceae =Scrophulariaceae s.l. (2001),recognition ofScrophulariaceae s.l.includ-
Scrophulariaceae Scrophulariaceae ing Buddlejaceae,Callitrichaceae,Globulariaceae,
Stilbaceae Stilbaceae Hippuridaceae, Myoporaceae and Plantaginaceae
Tetrachondraceae Tetrachondraceae
s.str. is not significantly less parsimonious than
=Pedaliaceae Trapellaceae
dividing them into the above five families, pro-
Verbenaceae Verbenaceae
vided that Calceolariaceae are excluded.This latter
family groups quite distant from Scrophulariaceae
s.l.Considering this and the comparatively small
Apart from the inclusion of Avicenniaceae in sample of genera analysed so far, treatment of
Acanthaceae here (Schwarzbach and McDade Scrophulariaceae in a wide circumscription
2002),in contrast to APG II,and the inclusion of is justifiable, but not when Buddlejaceae,
Trapellaceae in Pedaliaceae by APG II, also in Callitrichaceae, Globulariaceae, Hippuridaceae,
contrast to the present work,the major difference Myoporaceae and Plantaginaceae (and probably
between APG II and this FGVP volume lies in the Phrymaceae) are excluded, and Calceolaria
treatment of Scrophulariaceae s.l.and allies. and allies are included. However, it is very likely
Following the molecular evidence last summa- that eventually Calceolariaceae, Orobanchaceae,
rized and expanded by Olmstead et al.(2001),APG Paulowniaceae, Plantaginaceae/Veronicaceae
II divide Scrophulariaceae s.l. and allies into (incl.Callitrichaceae,Globulariaceae and Hippuri-
Orobanchaceae, Paulowniaceae, Plantaginaceae daceae), Schlegeliaceae, Scrophulariaceae s.str.
(incl.Callitrichaceae,Globulariaceae and Hippuri- (incl. Buddlejaceae and Myoporaceae) and an
daceae; these are the Veronicaceae of Olmstead expanded Phrymaceae will have to be recognized
et al. 2001), Schlegeliaceae and Scrophularia- because these lineages are, at least partly, more
ceae s.str. (incl. Buddlejaceae and Myoporaceae). closely related to other families of Lamiales
Calceolariaceae are recognized as an additional than to each other.Accordingly, the treatment of
family by Olmstead et al. (2001), and there is Scrophulariaceae s.l. (see Fischer, this Vol.)
increasing evidence that Phrymaceae, containing indicates the likely contents of the above lineages.
Phrymaand several genera mainly (but not only) As regards the treatment of Labiatae/
of Scrophulariaceae-Mimuleae, should be recog- Verbenaceae here and in APG II, in comparison
nized in an expanded circumscription (Beardsley with mainly Cronquist (1981) but partly also
and Olmstead 2002).In contrast,Scrophulariaceae Thorne (1992) and Takhtajan (1997),major parts
in the present treatment include Calceolariaceae, of Verbenaceae as traditionally circumscribed
Orobanchaceae, Paulowniaceae, and Schlegeli- have been transferred to Labiatae,based on sub-