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LPWG • Phylogeny and classification of the Leguminosae
A new subfamily classification of the Leguminosae based on
a taxonomically comprehensive phylogeny
The Legume Phylogeny Working Group (LPWG)
Recommended citation: LPWG (2017)
This paper is a product of the Legume Phylogeny Working Group, who discussed, debated and agreed on the classification of the
Leguminosae presented here, and are listed in alphabetical order. The text, keys and descriptions were written and compiled by a subset
of authors indicated by §. Newly generated matK sequences were provided by a subset of authors indicated by *. All listed authors
commented on and approved the final manuscript.
Nasim Azani,1 Marielle Babineau,2* C. Donovan Bailey,3* Hannah Banks,4 Ariane R. Barbosa,5* Rafael
Barbosa Pinto,6* James S. Boatwright,7* Leonardo M. Borges,8* Gillian K. Brown,9* Anne Bruneau,2§*
Elisa Candido,6* Domingos Cardoso,10§* Kuo-Fang Chung,11* Ruth P. Clark,4 Adilva de S. Conceição,12*
Michael Crisp,13* Paloma Cubas,14* Alfonso Delgado-Salinas,15 Kyle G. Dexter,16* Jeff J. Doyle,17 Jérôme
Duminil,18* Ashley N. Egan,19* Manuel de la Estrella,4§* Marcus J. Falcão,20 Dmitry A. Filatov,21* Ana
Paula Fortuna-Perez,22* Renée H. Fortunato,23 Edeline Gagnon,2* Peter Gasson,4 Juliana Gastaldello
Rando,24* Ana Maria Goulart de Azevedo Tozzi,6 Bee Gunn,13* David Harris,25 Elspeth Haston,25 Julie A.
Hawkins,26* Patrick S. Herendeen,27§ Colin E. Hughes,28§* João R.V. Iganci,29* Firouzeh Javadi,30* Sheku
Alfred Kanu,31 Shahrokh Kazempour-Osaloo,32* Geoffrey C. Kite,4 Bente B. Klitgaard,4§ Fábio J.
Kochanovski,6 Erik J.M. Koenen,28§* Lynsey Kovar,3* Matt Lavin,33* Marianne le Roux,34 Gwilym P. Lewis,4§
Haroldo C. de Lima,20 Maria Cristina López-Roberts,5* Barbara Mackinder,25§* Vitor Hugo Maia,35*
Valéry Malécot,36 Vidal F. Mansano,20* Brigitte Marazzi,37* Sawai Mattapha,26* Joseph T. Miller,38 Chika
Mitsuyuki,39* Tania Moura,40* Daniel J. Murphy,41 Madhugiri Nageswara-Rao,3* Bruno Nevado,21* Danilo
Neves,4* Dario I. Ojeda,18 R. Toby Pennington,25§* Darién E. Prado,42 Gerhard Prenner,4 Luciano Paganucci
de Queiroz,5§* Gustavo Ramos,10 Fabiana L. Ranzato Filardi,20* Pétala G. Ribeiro,5 María de Lourdes
Rico-Arce,4 Michael J. Sanderson,43 Juliana Santos-Silva,12* Wallace M.B. São-Mateus,44* Marcos J.S.
Silva,45* Marcelo F. Simon,46* Carole Sinou,2§* Cristiane Snak,5* Élvia R. de Souza,12* Janet Sprent,47 Kelly
P. Steele,48* Julia E. Steier,49* Royce Steeves,2* Charles H. Stirton,50 Shuichiro Tagane,39* Benjamin M.
Torke,51* Hironori Toyama,39* Daiane Trabuco da Cruz,5* Mohammad Vatanparast,19* Jan J. Wieringa,52§*
Michael Wink,53* Martin F. Wojciechowski,49§* Tetsukazu Yahara,39* Tingshuang Yi54 & Erin Zimmerman2
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
Department of Plant Science, University of Tehran, Iran
Institut de Recherche en Biologie Végétale and Département de Sciences Biologiques, Université de Montréal, Canada
Department of Biology, New Mexico State University, Las Cruces, U.S.A.
Royal Botanic Gardens, Kew, Richmond, Surrey, U.K.
Departamento Ciências Biológicas, Universidade Estadual de Feira de Santana, Brazil
Departamento de Biologia Vegetal, Universidade Estadual de Campinas, Brazil
Department of Biodiversity and Conservation Biology, University of the Western Cape, Cape Town, South Africa
Departamento de Botânica, Universidade Federal de São Carlos, Brazil
School of BioSciences, University of Melbourne, Australia and Queensland Herbarium, Toowong, Australia
Instituto de Biologia, Universidade Federal da Bahia, Salvador, Brazil
School of Forestry and Resource Conservation, National Taiwan University, Taiwan
Departamento de Educação, Campus VIII, Universidade do Estado da Bahia, Paulo Afonso, Brazil
Research School of Biology, The Australian National University, Acton, Australia
Departamento de Biología Vegetal II, Universidad Complutense, Madrid, Spain
Instituto de Biología – Botánica, Universidad Nacional Autónoma de México, México
School of GeoSciences, University of Edinburgh, U.K.
Plant Biology Department, Cornell University, Ithaca, New York, U.S.A.
Service Évolution Biologique et Écologie, Université Libre de Bruxelles, Belgium
Department of Botany, Smithsonian Institution, National Museum of Natural History, Washington D.C., U.S.A.
Instituto de Pesquisas, Jardim Botânico do Rio de Janeiro, Brazil
Department of Plant Sciences, University of Oxford, U.K.
Received: 6 Aug 2016 | returned for (first) revision: 2 Oct 2016 | (last) revision received: 19 Dec 2016 | accepted: 19 Dec 2016 || publication date(s):
online fast track, n/a; in print and online issues, 23 Feb 2017 || Published online “open-access” under the terms of the Creative Commons Attribution
4.0 (CC BY 4.0) License || © International Association for Plant Taxonomy (IAPT) 2017
44
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https://doi.org/10.12705/661.3
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LPWG • Phylogeny and classification of the Leguminosae
22
23
24
25
26
27
28
29
30
31
32
33
34
Departamento de Botânica, Universidade Estadual Paulista, Botucatu, São Paulo, Brazil
Instituto de Recursos Biológicos, CIRN-INTA, CONICET & University of Morón, Buenos Aires, Argentina
Ciências Ambientais, Universidade Federal do Oeste da Bahia, Barreiras, Brazil
Royal Botanic Gardens, Edinburgh, U.K.
School of Biological Sciences, University of Reading, U.K.
Chicago Botanic Garden, Glencoe, Illinois, U.S.A.
Department of Systematic and Evolutionary Botany, University of Zurich, Switzerland
Departamento de Botânica, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil
Institute of Decision Science for a Sustainable Society, Kyushu University, Fukuoka, Japan
Department of Agriculture and Animal Health, University of South Africa, South Africa
Department of Plant Biology, Tarbiat Modares University, Tehran, Iran
Plant Sciences & Plant Pathology, Montana State University, Bozeman, Montana, U.S.A.
South African National Biodiversity Institute, Silverton, South Africa and Department of Botany and Plant Biotechnology,
University of Johannesburg, South Africa
35 Departamento de Biologia, Pontifícia Universidade Católica do Rio de Janeiro, Brazil
36 Agrocampus-Ouest, INRA, Université d’Angers, France
37 Museo Cantonale di Storia Naturale, Lugano, Switzerland
38 Office of International Science and Engineering, National Science Foundation, Arlington, Virginia, U.S.A.
39 Department of Biology, Kyushu University, Fukuoka, Japan
40 Missouri Botanical Garden, Saint Louis, Missouri, U.S.A.
41 Plant Sciences and Biodiversity, Royal Botanic Gardens Victoria, Melbourne, Australia
42 Botánica, IICAR-CONICET, Facultad de Ciencias Agrarias, Universidad Nacional de Rosario, Zavalla, Argentina
43 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, U.S.A.
44 Sistemática e Evolução, Universidade Federal do Rio Grande do Norte, Brazil
45 Departamento de Botânica, Instituto de Ciências Biológicas, Universidade Federal de Goiás, Brazil
46 Embrapa Recursos Geneticos e Biotecnologia, Brasília, Brazil
47 Department of Plant Sciences, University of Dundee, U.K.
48 Faculty of Science and Mathematics, Arizona State University Polytechnic, Mesa, Arizona, U.S.A.
49 School of Life Sciences, Arizona State University, Tempe, Arizona, U.S.A.
50 Department of Biological Sciences, Bolus Herbarium, University of Cape Town, Rondebosch, South Africa
51 Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York, U.S.A.
52 Naturalis Biodiversity Center, Leiden, The Netherlands
53 Institute of Pharmacy and Molecular Biotechnology, Heidelberg University, Germany
54 Kunming Institute of Botany, Chinese Academy of Sciences, China
Author for correspondence: Anne Bruneau, anne.bruneau@umontreal.ca
DOI https://doi.org/10.12705/661.3
Abstract The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally
recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK gene
sequences, and including near-complete sampling of genera (698 of the currently recognised 765 genera) and ca. 20% (3696)
of known species. The matK gene region has been the most widely sequenced across the legumes, and in most legume lineages, this gene region is sufficiently variable to yield well-supported clades. This analysis resolves the same major clades as
in other phylogenies of whole plastid and nuclear gene sets (with much sparser taxon sampling). Our analysis improves upon
previous studies that have used large phylogenies of the Leguminosae for addressing evolutionary questions, because it maximises generic sampling and provides a phylogenetic tree that is based on a fully curated set of sequences that are vouchered
and taxonomically validated. The phylogenetic trees obtained and the underlying data are available to browse and download,
facilitating subsequent analyses that require evolutionary trees. Here we propose a new community-endorsed classification of
the family that reflects the phylogenetic structure that is consistently resolved and recognises six subfamilies in Leguminosae:
a recircumscribed Caesalpinioideae DC., Cercidoideae Legume Phylogeny Working Group (stat. nov.), Detarioideae Burmeist.,
Dialioideae Legume Phylogeny Working Group (stat. nov.), Duparquetioideae Legume Phylogeny Working Group (stat. nov.),
and Papilionoideae DC. The traditionally recognised subfamily Mimosoideae is a distinct clade nested within the recircumscribed Caesalpinioideae and is referred to informally as the mimosoid clade pending a forthcoming formal tribal and/or cladebased classification of the new Caesalpinioideae. We provide a key for subfamily identification, descriptions with diagnostic
charactertistics for the subfamilies, figures illustrating their floral and fruit diversity, and lists of genera by subfamily. This
new classification of Leguminosae represents a consensus view of the international legume systematics community; it invokes
both compromise and practicality of use.
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Keywords Caesalpinioideae; Cercidoideae; Detarioideae; Dialioideae; Duparquetioideae; mimosoid clade; Papilionoideae;
plastid matK phylogeny
Supplementary Material Electronic Supplement (Fig. S1), voucher information (Table S1), matK DNA sequence alignment
(Data File A), phylogenetic tree files (Data Files B–F) and a poster illustrating the new classification of the Leguminosae
(Figs. S2 & S3) are available in the Supplementary Data section of the online version of this article at http://www.
ingentaconnect.com/content/iapt/tax and on Data Dryad (DOI: https://doi.org/10.5061/dryad.61pd6).
From arctic circle to tropics, desert to pergola, bacteria to
plough, field to mouth, and legend to science, Leguminosae invest
our lives, and a feeble backwash seeps through our universities.
We wait that treatise which will quicken the herbarium into the
living tree of phylogeny.
Corner (1976: 162)
A general system of classification that is reasonably natural,
mnemonic and traditional is likely to be most useful for most purposes.
Polhill & al. (1981: 23–24)
Nutritious seeds for a sustainable future — The U.N. General
Assembly declared 2016 the International Year of Pulses to raise
awareness of the many benefits of legumes.
INTRODUCTION
The economically and ecologically important family
Leguminosae (Lewis & al., 2005; Yahara & al., 2013), or
Fabaceae (see Lewis & Schrire, 2003), has been the focus
of numerous recent phylogenetic analyses at the subfamily,
tribe and generic-group levels (see LPWG, 2013a and references therein). These, as well as phylogenies of the family as
a whole (Käss & Wink, 1996; Doyle & al., 1997, 2000; Kajita
& al., 2001; Wojciechowski & al., 2004; Lavin & al., 2005;
McMahon & Sanderson, 2006; Bruneau & al., 2008; Simon &
al., 2009; Cardoso & al., 2013b; LPWG, 2013a), all indicate that
the currently accepted classification of the family into the three
well-known, long-recognised and widely accepted subfamilies,
Caesalpinioideae DC., Mimosoideae DC., and Papilionoideae
DC., is outdated and does not reflect our current knowledge of
phylogenetic relationships in the family.
With close to 770 genera and over 19,500 species (Lewis &
al., 2005, 2013; LPWG, 2013a), the Leguminosae is the thirdlargest angiosperm family in terms of species numbers after
Asteraceae and Orchidaceae. Economically, Leguminosae is
second in importance only to Poaceae. It is estimated, for example, that total world exports of pulses (i.e., legume crops
harvested for their dry seeds) have more than doubled between
1990 and 2012, expanding from 6.6 to 13.4 million tons, and
in 2012 the value of pulse exports was estimated at US$ 9.5
billion (Food and Agriculture Organisation [FAO]: http://
www.fao.org/pulses-2016/en/). The United Nations General
Assembly designated 2016 the International Year of Pulses to
promote awareness of their nutritional benefits, importance in
food security and sustainable agriculture, and in mitigating
biodiversity loss and climate change. Legumes are important
food crops providing highly nutritious sources of protein and
micronutrients that can greatly benefit health and livelihoods,
46
particularly in developing countries (Yahara & al., 2013).
Legumes have been domesticated alongside grasses in different areas of the world since the beginnings of agriculture and
have played a key role in its early development (Gepts & al.,
2005; Hancock, 2012). Legumes are also uniquely important
as fodder and green manure in both temperate and tropical
regions, and are used for their wood, tannins, oils and resins,
in the manufacture of varnishes, paints, dyes and medicines,
and in the horticultural trade.
Legumes are cosmopolitan in distribution, representing
important ecological constituents in almost all biomes across the
globe and occur in even the most extreme habitats (Schrire & al.,
2005a, b). They constitute significant elements in terms of both
species diversity and abundance, in lowland wet tropical forests
in Africa, South America, and Asia (Yahara & al., 2013), and
they dominate dry forests and savannas throughout the tropics
(DRYFLOR, 2016), and also occur in Mediterranean, desert
and temperate regions, up to high latitudes and at high elevations. They can be large emergent tropical trees with buttresses,
small ephemeral annual herbs, climbing annuals or perennials
with tendrils, desert shrubs, geoxylic subshrubs, woody lianas
and, less commonly, aquatics. Flower symmetry spans the full
range from radially symmetric (actinomorphic) to bilaterally
symmetric (zygomorphic) and asymmetric flowers, which are
in turn adapted to a wide range of pollinators such as insects,
birds and bats. The ability of the majority of legume species
to fix atmospheric nitrogen in symbiosis with soil rhizobia is
perhaps the best-known ecological characteristic of the family;
however, not all legumes form associations with nitrogen-fixing
bacteria. Overall, the family is morphologically, physiologically
and ecologically exceptionally diverse, representing one of the
most spectacular examples of evolutionary diversification in
plants. All of these characteristics have led to a continued fascination with the biology, diversity and evolution of the family, the
evolution of functional traits, and the ecology and biogeography
of the family by legume biologists (e.g., Stirton & Zarucchi,
1989; Lavin & al., 2004; Schrire & al., 2005a, b; Sprent, 2007,
2009; Champagne & al., 2007; Simon & al., 2009; BouchenakKhelladi & al., 2010; Cannon & al., 2010, 2015; Pennington &
al., 2010; Doyle, 2011; Simon & Pennington, 2012; Koenen & al.,
2013; Oliveira-Filho & al., 2013; Moncrieff & al., 2014; Werner
& al., 2014, 2015; Dugas & al., 2015; BFG, 2015).
Here we propose a new subfamilial classification of the
family Leguminosae that takes into account the phylogenetic
pattern that is consistently resolved in numerous recent studies.
This new classification is proposed and endorsed by the legume
systematics community as reflected in the use of the Legume
Phylogeny Working Group (LPWG) as the authority for all
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new names proposed. The Legume Phylogeny Working Group
was established explicitly to develop and foster collaborative
research towards a comprehensive phylogeny and classification
for Leguminosae (LPWG, 2013a).
The new classification proposed here follows a traditional
Linnaean approach but is compatible with and complementary to emerging clade-based classifications of individual legume subfamilies (Wojciechowski, 2013). Rank-free naming
of clades within (and across) subfamilies is already well-established and increasingly prevalent in the legume literature
(e.g., Dalbergioid clade, Lavin & al., 2001; inverted repeat
[IR]-lacking clade, Wojciechowski & al., 2000; Umtiza clade,
Herendeen & al., 2003; Acacia s.l. clade, Miller & al., 2014),
and additional important clades will continue to be named even
after a fully fledged and stable subfamily and tribal classification is established. As noted by Wojciechowski (2013), use of
Linnaean names does not preclude a system that also defines
and names clades and their overall relationships outside of the
traditional Linnaean framework. Instead, the two are considered complementary and necessary for developing a stable,
flexible and useful classification of legumes.
THE NEW SUBFAMILY CLASSIFICATION
The monophyly of the family Leguminosae is strongly
supported in all molecular phylogenetic analyses, regardless
of taxon or gene sampling (see LPWG, 2013a and references
therein). Indeed, despite uncertainty over their closest relatives (cf. Dickison, 1981; APG III, 2009; Bello & al., 2009),
the monophyly and distinctiveness of the Leguminosae have
never been questioned in terms of morphology since the family
was first established (Adanson, 1763; Jussieu, 1789; Polhill
& Raven, 1981; Polhill, 1994; Lewis & al., 2005; Bello &
al., 2012). The most conspicuous characteristic of the family
is, with only a few exceptions, a single superior carpel with
one locule, marginal placentation, and usually two to many
ovules, in two alternating rows on a single placenta (Lewis &
al., 2005). However, legume systematists have been aware for
a long time of the discrepancy between the current subfamily
classification and emerging phylogenetic results (Irwin, 1981;
Käss & Wink, 1996; Doyle & al., 1997), most notably the long
known paraphyly of subfamily Caesalpinioideae, as well as
many other problematic issues, such as lack of monophyly of
many tribes and subtribes. This means that the phylogenetic
structure of the family is not directly reflected in the current
classification (Lewis & al., 2005). Thus, legume biologists
studying particular clades have invented and used informal
clade names that are biologically meaningful and appropriate
for their study questions. This has resulted in a proliferation of
informally named clades that can be inconsistent, ad hoc, and
sometimes contradictory across studies, and which can lead
to nomenclatural confusion unless they are properly defined
(LPWG, 2013a, b; Wojciechowski, 2013). This is important not
just within the legume taxonomic community but also for the
legume biology, genomics, and indeed the wider evolutionary
biology community as a whole (e.g., Cannon & al., 2015).
In contrast to some other large angiosperm families where
the subfamily rank is perhaps not as widely recognised or used
outside the immediate taxonomic community (e.g., Poaceae,
Grass Phylogeny Working Group, 2001, 2012; Asteraceae,
Panero & Funk, 2002, Funk & al., 2009), in legumes, the subfamily has always been a widely used and central rank. The
three currently recognised subfamilies have long been considered as distinct groups and have often been recognised at
the family rank (e.g., Hutchinson, 1964; Cronquist, 1981). In
1825, in his Prodromus, Candolle subdivided the Leguminosae
into four suborders (= subfamilies), naming for the first time
the three present-day subfamilies in addition to a fourth “suborder”, Swartzieae, now included in subfamily Papilionoideae.
This system was elaborated upon by Bentham (1865), who recognised three major groups within Leguminosae and whose
classification formed the basis for all subsequent classifications
of the family over the following 140 years (from, e.g., Taubert,
1891, to Polhill, 1994, and Lewis & al., 2005). In his Families
of flowering plants (1926) and Genera of flowering plants
(1964), Hutchinson raised the three subfamilies to the family
level, but grouped them in the order Leguminales, a system
that has been followed in a number of Floras (e.g., Hutchinson
& Dalziel, 1928; Görts-van Rijn, 1989; Orchard & Wilson,
1998–2001; Mori & al., 2002; see also Lewis & Schrire, 2003).
In the first volume of Advances in legume systematics (Polhill
& Raven, 1981), the three groups were recognised at the subfamily rank. Regardless of rank, these three groups have been
used as a division for identifying and classifying genera and
species in Floras and herbaria throughout the world since the
19th century. These groupings are taught in botany, floristics
and taxonomy courses, and are consistently used by agronomists, horticulturalists, and ecologists throughout the world. As
remarked by Polhill & al. (1981: 24), “the basic classification of
the family has remained remarkably stable and sensible. Users
of classifications provide a strong selective force […]”. Indeed,
although the generic membership of the three subfamilies has
changed somewhat over the centuries, these iconic groupings
have remained useful concepts for identifying this diverse
group of plants. Our objective here is to retain the utility of
these well-known groups as far as possible while at the same
time proposing a new classification that correctly reflects the
evolutionary relationships and emphasises the distinctive features of each of the subfamilies.
Despite tremendous progress in understanding phylogenetic relationships across the family (LPWG, 2013a), uncertainty remains regarding relationships amongst the six first
branching lineages of legumes and within certain clades
(Figs. 1 & 2) (Wojciechowski & al., 2004; Bruneau & al.,
2008; LPWG, 2013a). For example, relationships among early-branching papilionoids (Cardoso & al., 2012a, 2013b), the
large so-called Mimosoideae-Caesalpinieae-Cassieae clade,
or MCC clade sensu Doyle (2011, 2012) (Bruneau & al., 2008;
Manzanilla & Bruneau, 2012; ), and the Ingeae-Acacia s.str.
clade (Luckow & al., 2003; Simon & al., 2009) all lack resolution and support using conventional DNA sequence datasets
(i.e., a few kilobases of plastid DNA sequence data). However,
there is no uncertainty surrounding the paraphyly of subfamily
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Duparquetioideae
(1/1 genus, 1/1 species)
Cercidoideae
(12/12 genera, 96/ca. 335 species)
A.
Detarioideae
(79/84 genera, 327/ca. 760 species)
Dialioideae
(15/17 genera, 19/ca. 85 species)
Caesalpinioideae (incl. mimosoid clade)
(146/148 genera, 937/ca. 4400 species)
Papilionoideae
(445/503 genera, 2316/ca. 14,000 species)
B.
mimosoid clade
6.0
0.06
◄
Fig. 1. A, Bayesian consensus phylogenetic tree of 3842 matK sequences representing 3696 of the ca. 19,500 species and 698 of the 765 genera (Table
2) of Leguminosae (for 30 species, multiple varieties or subspecies were included) and 100 outgroup taxa (uncoloured) spanning core Eudicots
(see Appendix 1, Table S1). Branch lengths are proportional to numbers of matK substitutions. All subfamilies are supported with 1.0 posterior
probability (indicated as thicker lines) and 100% maximum likelihood bootstrap values (Fig. S1). Support is weak across the backbone of the grade
subtending the mimosoid clade, and this grade includes five or more lineages which would need to be recognised as additional small subfamilies if
Mimosoideae had been retained at a subfamilial rank. Duparquetioideae forms a polytomy with Cercidoideae, Detarioideae and the clade that groups
the other three subfamilies (but see Fig. 2, where Duparquetioideae is sister to the clade comprising Dialioideae, Caesalpinioideae and Papilionoideae based on analysis of a much larger plastid gene set). Numbers of genera and species (+ infraspecific taxa) sampled / currently recognised are
indicated for each subfamily. The phylogenetic tree can be visualised (e.g., with FigTree [http://tree.bio.ed.ac.uk/software/figtree/] or Dendroscope
[http://dendroscope.org/]; Huson & Scornavacca, 2012), and downloaded from Supplementary Data: Data file B. B, Schematic phylogeny based
on the matK Bayesian analysis showing the six subfamily classification of the Leguminosae, with clade sizes proportional to number of species.
A schematic figure illustrating the diversity of the six subfamilies is available for download as a poster (Figs. S2, S3).
48
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◄
Fig. 2. Phylogeny and subfamily classification of the Leguminosae, depicted on a 95% majority-rule Bayesian consensus tree based on analysis of
peptide sequences from 81 plastid encoded proteins, subsampling representative taxa from forthcoming phylogenomic analyses (E.J.M. Koenen
& al., in prep.). This analysis resolves the relationships of Duparquetioideae (cf. Fig. 1 based on analysis of matK alone). The tree is unresolved in
just a few places, including the root of the family and amongst clades in the Caesalpinioideae. All other nodes received 1.0 posterior probability,
except the two nodes marked with an asterisk, which have 0.99 posterior probability. The tree was inferred using PhyloBayes v.1.6j (Lartillot & al.,
2009) with the -CATGTR model selected and running two independent chains until they reached convergence. The six subfamilies are indicated
by the coloured boxes to the right of the phylogeny. Coloured branches indicate the three traditionally recognised subfamilies of Leguminosae: red
showing the paraphyletic old-sense Caesalpinioideae, blue the Mimosoideae and green the Papilionoideae.
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Caesalpinioideae and hence the need for a new subfamilial
classification (LPWG, 2013a, b). All adequately sampled phylogenetic analyses of the family indicate that the monophyletic
Mimosoideae and Papilionoideae are nested within a paraphyletic assemblage of caesalpinioid lineages. This is perhaps no
surprise. Already in 1981, in the preface to Advances in legume
systematics volume 1, based on morphology alone, H.S. Irwin
noted that Caesalpinioideae remained the most troublesome
segment of the family and that, inevitably, a greater number
of higher-level groups would need to be recognised.
The three traditional subfamilies were based essentially on a small set of conspicuous floral characters, particularly petal aestivation patterns (imbricate ascending in
Caesalpinioideae vs. imbricate descending in Papilionoideae
vs. valvate in Mimosoideae) and floral symmetry (variable
in Caesalpinioideae [Figs. 3–5]; radially symmetric [i.e., actinomorphic] in Mimosoideae [Fig. 6]; bilaterally symmetric [i.e., zygomorphic] in Papilionoideae [Figs. 7–9]). While
some of these floral characters may be useful for defining
Papilionoideae and Mimosoideae, they are extremely variable
across the traditional Caesalpinioidae (Tucker, 2003; Bruneau
& al., 2014), which cannot be defined or diagnosed based on
these characters. Furthermore, even for Papilionoideae and
Mimosoideae, most of these floral traits are now known to
be homoplasious (Pennington & al., 2000). For example,
individual species or clades marked by radially symmetric
flowers are independently derived multiple times across basal
Papilionoideae, a large assemblage of florally heterogeneous
lineages dominated by bilaterally symmetric flower morphology (Figs. 7–9) (Pennington & al., 2000; Cardoso & al., 2012b,
2013a; Ramos & al., 2016). Similarly, while Mimosoideae are
the most conspicuously biodiverse clade with radially symmetric flowers, other closely related lineages scattered across
the MCC clade also have radially symmetric, mimosoid-like
flowers (Fig. 5). Thus, despite the central importance of floral
characters in the traditional subfamilial classification, phylogenetic results over the past 20 years favour giving less weight
to floral morphology because it is so prone to evolutionary
modification and convergence, especially in the transition from
radial to bilateral floral symmetry, which can be achieved in
different ways.
There has been broad consensus about the need for a new
classification within the legume systematics community since
the first molecular phylogenies of the family were published
(Käss & Wink, 1996; Doyle & al., 1997). However, the multilineage paraphyletic structure of subfamily Caesalpinioideae
with respect to the monophyletic Mimosoideae and Papilionoideae poses significant questions about how many subfamilies
should be recognised. Furthermore, until recently, incomplete
sampling of many key genera in phylogenies suggested the need
for caution before establishing a new subfamilial classification.
More recent and densely sampled phylogenies (Luckow & al.,
2003; Wojciechowski & al., 2004; Lavin & al., 2005; Bruneau
& al., 2008; Simon & al., 2009; Cardoso & al., 2012a, 2013b),
as well as the matK phylogeny with its near-complete sampling
of genera that we present here (Figs. 1 & S1; Appendix 1), now
provide adequate taxon sampling and phylogenetic support to
50
reveal in sufficient and definitive detail the overall phylogenetic
structure of the family and allow us to properly evaluate the
options and arrive at the best solution for translating the phylogenetic tree into a new classification. Furthermore, the main
clades resolved in the matK phylogeny are also fully supported
in whole plastid genome sequence analyses (Fig. 2) (E.J.M.
Koenen & al., in prep.), and are corroborated by phylogenetic
analyses of orthologous nuclear genes derived from representative sampling of multiple transcriptomes of all subfamilies,
except Duparquetioideae (E.J.M. Koenen & al., in prep.).
A concerted effort to arrive at a new classification was
initiated at the 6th International Legume Conference in
Johannesburg, South Africa, in January 2013. Specifically,
there was general consensus that sufficient data, in terms of
taxon sampling and phylogenetic support, were available to
propose a new subfamilial classification for Leguminosae, and
there was universal agreement that the number of subfamilies needed to be increased (LPWG, 2013b). There was also
broad agreement that several caesalpinioid clades (Cercideae,
Detarieae, Duparquetia, Dialiineae s.l.) could be appropriately,
uncontroversially and usefully recognised as new subfamilies,
alongside Papilionoideae. The central problem for a new subfamilial classification, was how to deal with the large clade
that includes the “Umtiza clade” or “grade”, “Caesalpinia
Group clade”, “Cassia clade”, “Peltophorum clade”, scattered
Dimorphandra Group genera, and which has Mimosoideae
nested within it, i.e., the large MCC clade (sensu Doyle, 2011,
2012). Several participants suggested that the whole MCC clade
should be recognized as a single subfamily (making a total of
six subfamilies), but with the disadvantage that mimosoids, in
the traditional sense, would no longer be recognised as a subfamily, which made some legume systematists uncomfortable.
The alternative, whereby Mimosoideae is retained as a subfamily, entails recognition of six to eight (or more) additional
small subfamilies to account for the multiple lineages that make
up the large paraphyletic assemblage subtending mimosoids
(Figs. 1, 2). However, many recognised that although resolution and support across this grade remains relatively weak in
current phylogenies (Fig. 1; Bruneau & al., 2008; Manzanilla
& Bruneau, 2012), improved resolution and support from larger
datasets (e.g., Fig. 2; E.J.M. Koenen & al., in prep.) was not
alone going to solve the problem of 6 vs. 11 or more subfamilies.
These two main options for a new classification were summarised, the points of agreement noted, and the foundations for
furthering the discussion presented in LPWG (2013b).
The advantages and disadvantages of these two main
options for a new subfamily classification (6 vs. 11, or more
subfamilies) were specifically discussed and evaluated at a
subsequent legume systematics symposium, held during the
Latin American Botanical Congress in October 2014, in Bahia,
Brazil. A document was then drafted summarising the advantages and disadvantages and circulated to a LPWG electronic
discussion group with wide, international membership for further discussion and opinion. The comments received from this
draft were taken into account when developing the classification presented here, subfamily descriptions were discussed at a
legume morphology workshop in Botucatu, Brazil (November
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Fig. 3. A–F, Cercidoideae; G, Duparquetioideae; H–L, Dialioideae. A, Cercis siliquastrum; B, Bauhinia galpinii; C, Bauhinia divaricata; D, Piliostigma
thonningii; E, Griffonia physocarpa; F, Schnella cupreonitens; G, Duparquetia orchidacea; H, Zenia insignis; I, Apuleia leiocarpa; J, Poeppigia
procera; K, Distemonanthus benthamianus; L, Kalappia celebica. — Photos: A & B, Colin Hughes; C, Jonathan Amith; D, E & K, Xander van der
Burgt; F & I, Domingos Cardoso; G, Paul Hoekstra; H, Shijin Li; J, Luciano P. de Queiroz; L, Liam Trethowan.
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Fig. 4. Detarioideae. A, Goniorrhachis marginata; B, Hymenaea stigonocarpa; C, Daniellia ogea; D, Peltogyne chrysopis; E, Brodriguesia santosii;
F, Brownea longipedicellata; G, Amherstia nobilis; H, Brachycylix vageleri; I, Cryptosepalum tetraphyllum; J, Paramacrolobium coeruleum; K,
Gilbertiodendron quinquejugum; L, Aphanocalyx pteridophyllus. — Photos: A, D & F, Domingos Cardoso; B, Luciano P. de Queiroz; C, I, J & L,
Xander van der Burgt; E, Gwilym Lewis; G, Timothy Utteridge; H, Emilio Constantino; K, Jan Wieringa.
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Fig. 5. Caesalpinioideae I. A, Gleditsia amorphoides; B, Pterogyne nitens; C, Batesia floribunda; D, Moldenhawera blanchetiana; E, Cassia fistula;
F, Tachigali rugosa; G, Arapatiella psilophylla; H, Caesalpinia cassioides; I, Arquita grandiflora; J, Delonix floribunda; K, Campsiandra comosa;
L, Dimorphandra pennigera. — Photos: A, B, D, F & G, Domingos Cardoso; C & L, Projecto Flora Reserva Ducke, INPA/DFID, comm. Mike
Hopkins; E, Gwilym Lewis; H & I, Colin Hughes; J, David Du Puy; K, Luciano P. de Queiroz.
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Fig. 6. Caesalpinioideae II. A, Chidlowia sanguinea; B, Entada chrysostachys; C, Gagnebina commersoniana; D, Lemurodendron capuronii; E,
Neptunia plena; F, Mimosa benthamii; G, Acacia dealbata; H, Senegalia sakalava; I, Inga calantha; J, Inga grazielae; K, Macrosamanea amplissima; L, Albizia grandibracteata. — Photos: A, Xander van der Burgt; B–D, H, K & L, Erik Koenen; E–G, Colin Hughes; I, Flora do Acre, comm.
Rosangela Melo; J, Domingos Cardoso.
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Fig. 7. Papilionoideae I. A, Castanospermum australe; B, Petaladenium urceoliferum; C, Pterodon abruptus; D, Swartzia acutifolia; E, Trischidium
molle; F, Exostyles venusta; G, Harleyodendron unifoliolatum; H, Haplormosia monophylla; I, Ormosia lewisii; J, Harpalyce lanata; K, Leptolobium
brachystachyum; L, Camoensia brevicalyx. — Photos: A–G & I–K, Domingos Cardoso; H, Jan Wieringa; L, André van Proosdij.
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Fig. 8. Papilionoideae II. A, Uleanthus erythrinoides; B, Cadia purpurea; C, Sophora cf. microphylla; D, Virgilia divaricata; E, Cyclopia pubescens;
F, Lupinus weberbaueri; G, Dalea botterii; H, Errazurizia megacarpa; I, Zornia reticulata; J, Poiretia tetraphylla; K, Pterocarpus amazonum; L,
Baphia leptobotrys. — Photos: A, I & K, Domingos Cardoso; B, Wolfgang Stuppy; C, Gwilym Lewis; D & E, Stephen Boatwright; F, H & J, Colin
Hughes; G, Donovan Bailey; L, Jan Wieringa.
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Fig. 9. Papilionoideae III. A, Chorizema glycinifolium; B, Bossiaea walkeri; C, Mucuna gigantea; D, Chadsia longidentata; E, Canavalia brasiliensis; F, Erythrina velutina; G, Gliricidia robusta; H, Poissonia weberbaueri; I, Anthyllis montana; J, Astragalus uniflorus; K, Trifolium rubens; L,
Pisum sativum subsp. biflorum. — Photos: A & B, Michael Crisp; C, Timothy Utteridge; D, Erik Koenen; E, Domingos Cardoso; F, Luciano P. de
Queiroz; G & I–L, Colin Hughes; H, Justin Moat.
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Fig. 10. Legume fruit diversity I. A, Cercidoideae; B, Duparquetioideae; C, Dialioideae, D & E, Detarioideae; F– L, Caesalpinioideae. A, Griffonia
physocarpa; B, Duparquetia orchidacea; C, Dialium guianense; D, Brodriguesia santosii; E, Berlinia razzifera, held by Jean-Claude Mouzanda; F,
Eligmocarpus cynometroides; G, Heteroflorum sclerocarpum; H, Erythrostemon coccineus; I, Entada polystachya; J, Prosopis ferox; K, Mimosa
townsendii; L, Cojoba arborea. — Photos: A & B, Xander van der Burgt; C, Domingos Cardoso; D, G, H, J & L, Colin Hughes; E, David Harris;
F, Felix Forest; I & K, Gwilym Lewis.
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Fig. 11. Legume fruit diversity II. A & B, Caesalpinioideae; C–L, Papilionoideae. A, Abarema jupunba; B, Inga feuillei; C, Swartzia parvipetala; D,
Andira micrantha; E, Crotalaria cf. stipularia; F, Pterocarpus angolensis; G, Dalbergia lemurica; H, Machaerium millei; I, Carmichaelia cf. aligera;
J, Erythrina madagascariensis; K, Piscidia grandifolia; L, Phaseolus spp. — Photos: A & D, Projecto Flora Reserva Ducke, INPA/DFID, comm.
Mike Hopkins; B & I, Wolfgang Stuppy; C, James Ratter; E, F & H, Gwilym Lewis; G & J, David Du Puy; K & L, Colin Hughes.
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2015), and draft manuscripts circulated again to the LPWG
membership for further comment prior to submission of this
paper for publication.
After broad consultation within the legume systematics community, it was generally agreed that a six subfamily
classification is the most appropriate option for naming subfamilies in a Linnaean system (Figs. 1, 2, S2 & S3). The six
subfamily option is based on a set of clades with robust support (1.00 Bayesian posterior probabilities and 100% maximum likelihood bootstrap values in Figs. 1, 2 & S1) that are
each subtended by long branches: Cercidoideae (Fig. 3A–F),
Detarioideae (Fig. 4), Duparquetioideae (Fig. 3G), Dialioideae
(Fig. 3H–L), Papilionoideae (Figs. 7–9), and the recircumscribed
Caesalpinioideae (which equates to the MCC clade; Figs. 5 & 6).
In addition to the molecular support all six subfamilies have
support from morphological data (Table 1). While morphological circumscription of the six subfamilies is not entirely
straightforward given the complex and homoplasious nature of
most morphological characters (Table 1; see Taxonomy below),
it is certainly no more difficult or problematic than for the traditional three subfamilies, for which the supposed diagnostic
morphological (mainly conspicuous floral) characters are beset
by numerous exceptions, and where Caesalpinioideae, as traditionally circumscribed, lacks obvious diagnosability. Although
Papilionoideae and the re-circumscribed Caesalpinioideae are
still large and heterogeneous clades, the former retains its current definition and generic membership (Polhill, 1994; Lewis &
al., 2005) (Table 2), while the latter is now more homogeneous,
including, for example, all legumes with bipinnate leaves and
most with extrafloral nectaries on the petiole and rachis (Fig. 2;
Cercidoideae
Detarioideae
Duparquetioideae
Caesalpinioideae
Papilionoideae
Trees, shrubs or
lianas, many with
tendrils, mostly unarmed but frequently
with prickles or
infrastipular spines;
branches rarely
modified into
cladodes
Usually unarmed
trees, sometimes
shrubs, rarely
suffruticose
Unarmed scrambling Unarmed trees
or shrubs, rarely
liana
suffruticose
Trees, shrubs, lianas,
suffruticose or functionally herbaceous,
unarmed or commonly armed with
prickles or spines
Usually unarmed
trees, shrubs, lianas,
herbs, or twining
vines with tendrils
Mostly lacking
Often present on
the underside, rarely
on the margins of
leaflets or on the
leaf rachis
Lacking
Lacking
Often present on
the petiole and / or
on the primary
and secondary
rachises, usually
between pinnae or
leaflet pairs, sometimes on stipules or
bracts
Lacking on petiole
and leaf rachis; occasionally present
on stipules, stipels,
bracts, or swollen
and nectar-secreting peduncles or
sepals
Lateral, free
Intrapetiolar
(i.e., somewhere
between the petiole
and the axillary
bud) and then free,
valvate and connected by chaffy
hairs, or fused,
either partly (only
at base) or entirely,
rarely lateral and
free
Lateral, free
Lateral, free or
absent
Lateral, free or
absent
Lateral, free or
absent, very rarely
interpetiolar
Unifoliolate or
bifoliolate
Usually paripinnate or bifoliolate,
rarely unifoliolate
Imparipinnate
Usually imparipinnate, rarely paripinnate, 1-foliolate
or palmately
compound
Commonly bipinnate, otherwise
pinnate, and then
mostly paripinnate,
rarely imparipinnate
or bifoliolate, modified into phyllodes
or lacking
Mostly pari- or
imparipinnate
or palmately
compound, commonly unifoliolate,
trifoliolate, rarely
bifoliolate or
tetrafoliolate
Leaves
Stipules
Specialised
extrafloral nectaries
Habit
Table 1. Comparative morphology, chemistry and chromosome numbers of the six subfamilies of Leguminosae. The text in bold highlights characters and character states that are particularly valuable for identifying members of the subfamilies. See glossary in Appendix 2 and Figs. 12 &
13 for definitions and illustrations of technical terms.
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Dialioideae
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Table 1) (e.g., Marazzi & al., 2012). The six subfamilies have
similar stem ages, all having apparently diverged soon after
the first appearance of the family (Lavin & al., 2005; Bruneau
& al., 2008; Simon & al., 2009).
The major disadvantage of adopting a six subfamily classification, namely abandoning the well-known Mimosoideae, is
mitigated by continuing to recognise this lineage as a distinct
clade, informally referred to as the mimosoid clade at this point,
but with scope to be formally named as a tribe within a new
Linnaean tribal classification, and/or in a rank-free clade-based
phylogenetic classification of new sense Caesalpinioideae, once
relationships within this subfamily are better resolved. It is
also worth noting that options recognising fewer than six subfamilies would both reduce morphological diagnosability and
result in subfamilies with even more unwieldy morphological
heterogeneity. The six subfamily option minimises the number of new Linnaean names, which is likely to be more easily
accepted by a wider user community, and we considered this
option as more likely to remain stable through time. With a
six subfamily system, we are ensuring greater nomenclatural
stability than a system that would describe 11 or more new subfamilies, particularly as several of the additional subfamilies
that would need to be recognised lack robust support in current
phylogenies being subtended by short branches (Figs. 1, 2 &
S1) (Bruneau & al., 2008; Manzanilla & Bruneau, 2012; E.J.M.
Koenen & al., in prep.) and might later need to be changed.
Although Caesalpinioideae DC. and Mimosoideae
DC. have equal priority under the International Code of
Nomenclature for algae, fungi and plants (Melbourne Code)
(McNeill & al., 2012) because they were published in the same
Cercidoideae
Detarioideae
Duparquetioideae
Dialioideae
Caesalpinioideae
Papilionoideae
Opposite (when
bifoliolate); blade
(when unifoliolate)
entire or bilobed
Opposite or alternate; translucent
glands sometimes
present
Opposite; blade
entire
Alternate, rarely
opposite
Mostly opposite,
rarely alternate
Opposite or alternate, sometimes
modified into
tendrils, rarely in
phyllodes
Raceme or
pseudoraceme
Raceme or panicle
Terminal raceme
Branched, thyrsoid
inflorescences, less
commonly racemes
with distichous
flower arrangement,
or flowers solitary
Globose, spikes,
panicles, racemes or
flowers in fascicles
Mostly racemes,
pseudoracemes or
panicles, less often
cymes, spicate or
capitate, or flowers
solitary
Large or minute
Large or small,
frequently petaloid,
valvate, imbricate
or partially fused
or partly fused
with the hypanthium, partially or
completely enclosing the bud
Small
Small or absent
Small or absent
Mostly small, rarely
large, valvate,
enveloping the bud
Bisexual, rarely
unisexual, slightly to
strongly bilaterally
symmetrical, sometimes papilionate
Bisexual or with
both bisexual and
male flowers, radially or slightly to
strongly bilaterally
symmetrical, but
never papilionate
Bisexual, strongly
bilaterally symmetrical, never
papilionate
Bisexual, radially or
slightly to strongly
bilaterally symmetrical, sometimes
papilionate
Usually bisexual,
rarely unisexual,
or bisexual flowers
combined with unisexual and / or sterile
flowers in heteromorphic inflorescences; radially, less
frequently bilaterally
symmetrical, sometimes papilionate or
asymmetric
Bisexual, rarely
unisexual, usually
bilaterally symmetrical, usually
papilionate, rarely
asymmetrical, radially symmetrical or
nearly so
Present, greatly
elongated to almost
absent
Present, elongated to
almost absent
Absent
Usually absent,
rarely present,
receptacle may be
broad and flattened,
bearing nectary-like
bodies
Lacking or cupular,
rarely tubular
Present or absent
Hypanthium
Flowers
Bracteoles
Inflorescence
Leaflets and
pinnae
Table 1. Continued.
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Detarioideae
Duparquetioideae
United in a spathaceous or 2–5-lobed
calyx or sepals free
Commonly 5 or 4
(two adaxial sepals
often fused), rarely
some or all absent or
more (–7)
4, unequal, free, the (3 or 4)–5–(6), free,
abaxial and adaxial equal to sub-equal
sepals cucullate
and sepaloid, the
laterals petaloid
5, rarely 2, 6 or
absent, free, when
present imbricate, the adaxial
petal innermost
and frequently
differentiated
0–5(–7), free, when
present imbricate,
the adaxial petal
generally outermost, all equal or
the adaxial large and
either the other 4
or only the abaxial
ones smaller to
rudimentary
5, free, imbricate,
the adaxial petal
outermost, adaxial
and two lateral
petals ovate, two
abaxial petals straplike, oblong, all 5
petals with stalked
glands along their
margins
5 or fewer (0, 1, 3,
4), rarely 6, free, imbricate, the adaxial
petal innermost, all
equal to sub-equal
(3–)5(–6), free or
fused, or petal whorl
lacking, valvate
or imbricate,
then adaxial petal
innermost
Usually (0–)5(–6),
rarely 1 (standard)
petal and 4 absent,
imbricate, the
adaxial petal
outermost, in radially symmetrical
flowered species,
corolla with 5
small or undifferentiated petals,
less often only one
(standard) petal
is present or all
petals absent
Usually 10 (sometimes fewer) in two
whorls of alternate
length
Usually 10, sometimes 2–numerous
4
5 or fewer, rarely
6–10, uniform,
rarely dimorphic
Diplostemonous
or haplostemonous, sometimes
reduced to 3, 4 or
5, frequently many
(100+), sometimes
heteromorphic,
some or all sometimes modified or
staminodial
Usually 10, rarely 9
or many
Filaments partly
connate or free
Filaments partly
connate or free
Filaments free
Filaments free
Filaments free or
connate
Filaments usually
connate into a
sheath or tube,
uppermost filament
wholly or partly
free, sometimes all
filaments free
Mostly uniform,
dorsifixed, dehiscing
via longitudinal slits
or central pores;
reduced stamens or
staminodes sometimes present
Mostly uniform,
dorsifixed or basifixed, dehiscing via
longitudinal slits
Uniform, basifixed,
with pointed appendages, dehiscing
via short apical,
poricidal slits; anthers post-genitally
fused into a curving
synandrium
Uniform, rarely dimorphic, basifixed,
rarely dorsifixed,
dehiscing via longitudinal slits, often
reduced to short apical, poricidal slits;
staminodes present
or absent
Uniform or heteromorphic, basifixed
or dorsifixed, often
with a stipitate or
sessile apical gland,
dehiscing via longitudinal slits or apical
or basal poricidal
slits or pores
Uniform or dimorphic, basifixed or
dorsifixed, dehiscing via longitudinal
slits
Pollen
Monads, 3-colporate, 3–6-colpate,
3-porate, 3-pororate,
3–4-colporoidate or
inaperturate, rarely
in tetrads
Monads, mostly
3-colporate with
a vast array of
sculptures
Monads, asymmetrical, one equatorial-encircling
ectoaperture with
two equatorial
endoapertures
Monads tricolporate, with punctate
or finely reticulate,
rarely striate sculpture patterns
Monads, tricolporate
or porate tetrads,
bitetrads or
polyads, sculpture
pattern never striate
Monads, mostly
3-colporate,
3-colpate or
3-porate
1-carpellate, stipitate, stipe free or
adnate to the wall of
the hypanthium
1-carpellate, stipitate, stipe free or
adnate to the wall of
the hypanthium
1-carpellate, stipitate, stipe free
1-carpellate, sometimes 2-carpellate,
stipitate or sessile,
stipe free
Usually 1-carpellate,
rarely polycarpellate, stipitate or
sessile, stipe free
Usually 1-carpellate, rarely 2-carpellate, stipitate or
sessile, stipe free
Anthers
Stamen fusion
Stamens
Petals
Sepals
Cercidoideae
Gynoecium
Table 1. Continued.
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Papilionoideae
(3–)5, united at
(3–)5(–6), free or
fused, or sepal whorl least at the base,
sometimes entire
lacking
and splitting into
irregular lobes or
lobes dimorphic
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Detarioideae
Duparquetioideae
Dialioideae
Caesalpinioideae
Papilionoideae
Ovary
1–many-ovulate
Ovary
1–many-ovulate
Ovary 2–5-ovulate
Ovary frequently
2-ovulate, rarely
1–many-ovulate
Ovary
1–many-ovulate
Ovary
1–many-ovulate
Dehiscent pods,
often explosive with
twisted valves, or
indehiscent, then
generally samaroid
Mostly woody, dehiscent pods, sometimes indehiscent
and woody or thin
valved samaroid,
rarely filled with
pulpy mesocarp or
endocarp
Woody dehiscent
pods, 4-angled,
valves spirally
coiled
Commonly indehiscent drupaceous
or samaroid, rarely
dehiscent or the drupaceous fruit with
indurating endocarp
into one-seeded
segments
Commonly thinvalved, 1–manyseeded pod, dehiscent along one or
both sutures, also
often a lomentum,
a craspedium, or
thick and woody
and then indehiscent
or explosively
dehiscent, often
curved or spirally
coiled
Dehiscent pods
along one or both
sutures, or indehiscent, or loments,
samaras or drupes
With apical crescent-shaped hilum,
rarely circular; lens
inconspicuous, lacking pleurograms,
pseudopleurograms, wing or aril
Often overgrown,
sometimes hard and
then occasionally
with pseudopleurograms; occasionally
arillate
2–5, oblong to
ovoid, the testa
thick, lacking
pleurograms
1–2, rarely
more, lacking
pleurograms
Usually with an
open or closed
pleurogram
on both faces,
sometimes with
a fleshy aril or
sarcotesta, sometimes winged;
hilum usually apical; lens usually
inconspicuous
Usually with hard
testa, rarely overgrown, sometimes
with a fleshy aril
or sarcotesta; complex hilar valve,
elongate hilum and
lens usually present, pleurogram
lacking
Straight, very rarely
curved
Straight
Straight
Straight
Straight
Usually curved,
rarely straight
Vestured pits
in 2° xylem
Lacking
Present
Lacking
Usually lacking,
rarely present
Present
Present
Absent
Absent
Absent
Variably present and
indeterminate
Usually present,
either indeterminate
or determinate
2n = 14, 24, 26, 28
(42, 56)
2n mostly 24 (occasionally 16, 20, 22,
36, 68)
Unknown
2n = 28 (most genera not surveyed)
2n mostly 24, 26, 28
(but 14, 16, 52, 54,
56 also reported)
2n mostly 16, 18,
20, 22 (but 12,
14, 24, 26, 28,
30, 32, 38, 40,
48, 64, 84 also
reported)
Coumarins and
cyanogenic glucosides reported;
non-protein amino
acids common
(5-hydroxy-L-tryptophan only reported
to this subfamily)
Coumarins reported,
frequently with
terpenes (resins) and
non-protein amino
acids
Chemical characteristics unknown
Chemical characteristics unknown
Non-protein amino
acids frequently
reported; coumarins,
cyanogenic glucosids, phenylethylamine, tryptamines,
and β-carboline
alkaloids also
reported
Isoflavonoids,
prenylated
flavonoids, indolizidine or quinolizidine alkaloids
reported. Nonprotein amino
acids widespread,
some exclusively
found in the
subfamily (e.g.,
canavanine)
Chemistry
Chromosome
counts
Embyro
Seeds
Fruits
Ovules
Cercidoideae
Root
nodules
Table 1. Continued.
Absent
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Table 2. Genera of Leguminosae listed in alphabetical order within subfamilies. Recently synonymised genera are listed after the list of currently
recognised genera in each subfamily. Genera that have not been sampled in the matK phylogenetic analysis are identified by *. Genera of the
mimosoid clade in Caesalpinioideae are underlined.
CERCIDOIDEAE (12 genera, ca. 335 species): Adenolobus (Harv. ex Benth. & Hook.f.) Torre & Hillc.; Barklya F.Muell.; Bauhinia L.; Brenierea Humbert; Cercis L.; Gigasiphon Drake; Griffonia Baill.; Lysiphyllum (Benth.) de Wit; Phanera Lour.; Piliostigma Hochst.; Schnella
Raddi; Tylosema (Schweinf.) Torre & Hillc.
Recent synonym: Lasiobema (Korth.) Miq. = Phanera Lour.
DETARIOIDEAE (84 genera, ca. 760 species): Afzelia Sm.; Amherstia Wall.; Annea Mackinder & Wieringa; Anthonotha P.Beauv.; Aphanocalyx Oliver; Augouardia Pellegr.; Baikiaea Benth.; Barnebydendron J.H.Kirkbr.; Berlinia Sol. ex Hook. f.; Bikinia Wieringa; *Brachycylix
(Harms) R.S.Cowan; Brachystegia Benth.; Brandzeia Baill.; Brodriguesia R.S.Cowan; Brownea Jacq.; Browneopsis Huber; Colophospermum
J.Kirk ex J.Léonard; Copaifera L.; Crudia Schreb.; Cryptosepalum Benth.; Cynometra L.; Daniellia Benn.; Detarium Juss.; Dicymbe Spruce ex
Benth.; Didelotia Baill.; Ecuadendron D.A.Neill; Elizabetha Schomb. ex Benth.; Endertia Steenis & de Wit; Englerodendron Harms; Eperua
Aubl.; Eurypetalum Harms; Gabonius Wieringa & Mackinder; Gilbertiodendron J.Léonard; Gilletiodendron Vermoesen; Goniorrhachis Taub.;
Gossweilerodendron Harms; Guibourtia Benn.; Hardwickia Roxb.; Heterostemon Desf.; Humboldtia J.Vahl; Hylodendron Taub.; Hymenaea L.;
Hymenostegia (Benth.) Harms; Icuria Wieringa; Intsia Thouars; Isoberlinia Craib & Stapf; Isomacrolobium Aubrév. & Pellegr.; Julbernardia
Pellegr.; Kingiodendron Harms; Lebruniodendron J.Léonard; Leonardoxa Aubrév.; *Leucostegane Prain; Librevillea Hoyle; Loesenera Harms;
Lysidice Hance; Macrolobium Schreb.; Maniltoa Scheff.; *Michelsonia Hauman; *Micklethwaitia G.P.Lewis & Schrire; Microberlinia A.Chev.;
Neoapaloxylon Rauschert; Neochevalierodendron J.Léonard; Normandiodendron J.Léonard; Oddoniodendron De Wild.; Oxystigma Harms;
Paloue Aubl.; Paloveopsis R.S.Cowan; Paramacrolobium J.Léonard; Peltogyne Vogel; Plagiosiphon Harms; Polystemonanthus Harms; Prioria
Griseb.; *Pseudomacrolobium Hauman; Saraca L.; Schotia Jacq.; Scorodophloeus Harms; Sindora Miq.; Sindoropsis J.Léonard; Stemonocoleus
Harms; Talbotiella Baker f.; Tamarindus L.; Tessmannia Harms; Tetraberlinia (Harms) Hauman; Zenkerella Taub.
Recent synonym: Pellegriniodendron J.Léonard = Gilbertiodendron J.Léonard
DUPARQUETIOIDEAE (1 genus, 1 species): Duparquetia Baill.
DIALIOIDEAE (17 genera, ca. 85 species): *Androcalymma Dwyer; Apuleia Mart.; Baudouinia Baill.; Dialium L.; Dicorynia Benth.; Distemonanthus Benth.; Eligmocarpus Capuron; Kalappia Kosterm.; Koompassia Maingay ex Benth.; Labichea Gaudich. ex DC.; Martiodendron
Gleason; Mendoravia Capuron; Petalostylis R.Br.; Poeppigia C.Presl; Storckiella Seem.; *Uittienia Steenis; Zenia Chun
CAESALPINIOIDEAE (148 genera, ca. 4400 species; includes genera of the mimosoid clade, which are underlined): Abarema Pittier; Acacia
Mill.; Acaciella Britton & Rose; Acrocarpus Wight & Arn.; Adenanthera L.; Adenopodia C.Presl; Afrocalliandra E.R.Souza & L.P.Queiroz;
Alantsilodendron Villiers; Albizia Durazz.; Amblygonocarpus Harms; Anadenanthera Speg.; Arapatiella Rizzini & A.Mattos; Archidendron
F.Muell.; Archidendropsis I.C.Nielsen; Arcoa Urb.; Arquita E.Gagnon, G.P.Lewis & C.E.Hughes; Aubrevillea Pellegr.; Balizia Barneby &
J.W.Grimes; Balsamocarpon Clos; Batesia Spruce ex Benth. & Hook. f.; Biancaea Tod.; Blanchetiodendron Barneby & J.W.Grimes; Burkea
Benth.; Bussea Harms; Caesalpinia L.; Calliandra Benth.; Calliandropsis H.M.Hern. & P.Guinet; Calpocalyx Harms; Campsiandra Benth.;
Cassia L.; Cathormion Hassk.; Cedrelinga Ducke; Cenostigma Tul.; Ceratonia L.; Chamaecrista Moench; Chidlowia Hoyle; Chloroleucon
(Benth.) Britton & Rose; Cojoba Britton & Rose; Colvillea Bojer ex Hook.; Conzattia Rose; Cordeauxia Hemsl.; Coulteria Kunth; Cylicodiscus
Harms; Delonix Raf.; Denisophytum R.Vig.; Desmanthus Willd.; Dichrostachys (DC.) Wight & Arn.; Dimorphandra Schott; Dinizia Ducke;
Diptychandra Tul.; Ebenopsis Britton & Rose; Elephantorrhiza Benth.; Entada Adans.; Enterolobium Mart.; Erythrophleum Afzel. ex R.Br.;
Erythrostemon Klotzsch; Faidherbia A.Chev.; Falcataria (I.C.Nielsen) Barneby & J.W.Grimes; Fillaeopsis Harms; Gagnebina Neck. ex DC.;
Gelrebia E.Gagnon & G.P.Lewis; Gleditsia L.; Guilandina L.; Gymnocladus Lam.; Haematoxylum L.; Havardia Small; Hererolandia E.Gagnon &
G.P.Lewis; Hesperalbizia Barneby & J.W.Grimes; Heteroflorum M.Sousa; Hoffmannseggia Cav.; *Hultholia E.Gagnon & G.P.Lewis; Hydrochorea
Barneby & J.W.Grimes; *Indopiptadenia Brenan; Inga Mill.; Jacqueshuberia Ducke; Kanaloa Lorence & K.R.Wood; Lemurodendron Villiers;
Lemuropisum H.Perrier; Leucaena Benth.; Leucochloron Barneby & J.W.Grimes; Libidibia (DC.) Schltdl.; Lophocarpinia Burkart; Lysiloma
Benth.; Macrosamanea Britton & Rose ex Britton & Killip; Mariosousa Seigler & Ebinger; Melanoxylon Schott; Mezoneuron Desf.; Microlobius
C.Presl; Mimosa L.; Mimozyganthus Burkart; Moldenhawera Schrad.; Mora Schomb. ex Benth.; Moullava Adans.; Neptunia Lour.; Newtonia
Baill.; Pachyelasma Harms; Painteria Britton & Rose; Parapiptadenia Brenan; Pararchidendron I.C.Nielsen; Paraserianthes I.C.Nielsen; Parkia
R.Br.; Parkinsonia L.; Paubrasilia E.Gagnon, H.C.Lima & G.P.Lewis; Peltophorum (Vogel) Benth.; Pentaclethra Benth.; Piptadenia Benth.;
Piptadeniastrum Brenan; Piptadeniopsis Burkart; Pithecellobium Mart.; Pityrocarpa (Benth.) Britton & Rose; Plathymenia Benth.; Pomaria
Cav.; Prosopidastrum Burkart; Prosopis L.; Pseudopiptadenia Rauschert; Pseudoprosopis Harms; Pseudosamanea Harms; Pterogyne Tul.;
Pterolobium R.Br. ex Wight & Arn.; Recordoxylon Ducke; Samanea (Benth.) Merr.; Sanjappa E.R.Souza & M.V.Krishnaraj; Schizolobium Vogel;
Schleinitzia Warb. ex Nevling & Niezgoda; Senegalia Raf.; Senna Mill.; Serianthes Benth.; Sphinga Barneby & J.W.Grimes; Stachyothyrsus
Harms; Stenodrepanum Harms; Stryphnodendron Mart.; Stuhlmannia Taub.; Sympetalandra Stapf; Tachigali Aubl.; Tara Molina; Tetrapleura
Benth.; Tetrapterocarpon Humbert; Thailentadopsis Kosterm.; Umtiza Sim; Vachellia Wight & Arn.; Viguieranthus Villiers; Vouacapoua Aubl.;
Wallaceodendron Koord.; Xerocladia Harv.; Xylia Benth.; Zapoteca H.M.Hern.; Zuccagnia Cav.; Zygia P.Browne.
Recent synonyms: Guinetia L.Rico & M.Sousa = Calliandra Benth.; Marmaroxylon Killip = Zygia P.Browne; Poincianella Britton & Rose (in
part, including type) = Erythrostemon Klotzsch and (in part) = Cenostigma Tul.; Stahlia Bello = Libidibia (DC.) Schltdl.
PAPILIONOIDEAE (503 genera, ca. 14,000 species): Abrus Adans.; Acmispon Raf.; Acosmium Schott; Adenocarpus DC.; Adenodolichos
Harms; Adesmia DC.; Aenictophyton A.T.Lee; Aeschynomene L.; Afgekia Craib; Aganope Miq.; Airyantha Brummitt; Akschindlium H.Ohashi;
Aldina Endl.; Alexa Moq.; Alhagi Gagnebin; Alistilus N.E.Br.; Almaleea Crisp & P.H.Weston; Alysicarpus Desv.; Amburana Schwacke & Taub.;
Amicia Kunth; Ammodendron Fisch. ex DC.; Ammopiptanthus S.H.Cheng; Ammothamnus Bunge; Amorpha L.; Amphicarpaea Elliott ex Nutt.;
Amphimas Pierre ex Harms; Amphiodon Huber; Amphithalea Eckl. & Zeyh.; Anagyris L.; Anarthrophyllum Benth.; Ancistrotropis A.Delgado;
Andira Lam.; Angylocalyx Taub.; Antheroporum Gagnep.; Anthyllis L.; *Antopetitia A.Rich.; Aotus Sm.; Aphyllodium (DC.) Gagnep.; Apios
Fabr.; Apoplanesia C.Presl; Apurimacia Harms; Arachis L.; Argyrocytisus (Maire) Raynaud; Argyrolobium Eckl. & Zeyh.; Arthroclianthus
Baill.; Aspalathus L.; Astragalus L.; Ateleia (Moç & Sessé ex DC.) Benth.; Austrodolichos Verdc.; Austrosteenisia R.Geesink; Baphia Afzel.
& Lodd.; Baphiastrum Harms; Baphiopsis Benth. ex Baker; Baptisia Vent.; *Barbieria DC.; Behaimia Griseb.; Bionia Mart. ex Benth.; *Biserrula L.; Bituminaria Heist. ex Fabr.; Bobgunnia J.H.Kirkbr. & Wiersema; Bocoa Aubl.; Bolusafra Kuntze; Bolusanthus Harms; Bolusia Benth.;
Bossiaea Vent.; Bowdichia Kunth; Bowringia Champ. ex Benth.; Brongniartia Kunth; Brya P.Browne; Bryaspis P.A.Duvign.; *Burkilliodendron
Sastry; Butea Roxb. ex Willd.; Cadia Forssk.; Cajanus DC.; Calicotome Link; Callerya Endl.; Callistachys Vent.; Calobota Eckl. & Zeyh.;
Calophaca Fisch. ex DC.; Calopogonium Desv.; Calpurnia E.Mey.; Camoensia Welw. ex Benth.; Camptosema Hook. & Arn.; Campylotropis
Bunge; Canavalia DC.; Candolleodendron R.S.Cowan; Caragana Fabr.; Carmichaelia R.Br.; Carrissoa Baker f.; Cascaronia Griseb.;
64
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Table 2. Continued.
Castanospermum A.Cunn. ex Hook.; Centrolobium Mart. ex Benth.; Centrosema (DC.) Benth.; Chadsia Bojer; Chaetocalyx DC.; Chapmannia
Torr. & A. Gray; Chesneya Lindl. ex Endl.; Chorizema Labill.; Christia Moench; *Chrysoscias E.Mey.; Cicer L.; Cladrastis Raf.; Clathrotropis
(Benth.) Harms; Cleobulia Mart. ex Benth.; Clianthus Sol. ex Lindl.; Clitoria L.; Clitoriopsis R.Wilczek; *Cochlianthus Benth.; Cochliasanthus
Trew; Codariocalyx Hassk.; Collaea DC.; Cologania Kunth; Colutea L.; Condylostylis Piper; Cordyla Lour.; Corethrodendron Fisch. ex Basiner;
Coronilla L.; Coursetia DC.; Craibia Harms & Dunn; Cranocarpus Benth.; Craspedolobium Harms; Cratylia Mart. ex Benth.; *Cristonia
J.H.Ross; Crotalaria L.; Cruddasia Prain; Cullen Medik.; Cyamopsis DC.; Cyathostegia (Benth.) Schery; Cyclocarpa Afzel. ex Urb.; Cyclolobium Benth.; Cyclopia Vent.; Cymbosema Benth.; Cytisophyllum O.Lang; *Cytisopsis Jaub. & Spach; Cytisus Desf.; Dahlstedtia Malme;
Dalbergia L.f.; Dalbergiella Baker f.; Dalea Lucanus; Dalhousiea Wall. ex Benth.; Daviesia Sm.; Decorsea R.Vig.; Deguelia Aubl.; Dendrolobium (Wight & Arn.) Benth.; Dermatophyllum Scheele; Derris Lour.; *Desmodiastrum (Prain) A.Pramanik & Thoth.; Desmodium Desv.;
Dewevrea Micheli; Dichilus DC.; Dicraeopetalum Harms; Dillwynia Sm.; Dioclea Kunth; Diphyllarium Gagnep.; Diphysa Jacq.; Diplotropis
Benth.; Dipogon Liebm.; Dipteryx Schreb.; Discolobium Benth.; Disynstemon R.Vig.; Dolichopsis Hassl.; Dolichos L.; Dorycnium Mill.;
*Dorycnopsis Boiss.; Droogmansia De Wild.; Dumasia DC.; Dunbaria Wight & Arn.; Dussia Krug & Urb. ex Taub.; Dysolobium (Benth.)
Prain; Ebenus L.; *Echinospartum (Spach) Rothm.; Eleiotis DC.; *Eminia Taub.; Endosamara R.Geesink; Eremosparton Fisch. & C.A.Mey.;
Erichsenia Hemsl.; Erinacea Adans.; Eriosema (DC.) Desv.; Erophaca Boiss.; Errazurizia Phil.; Erythrina L.; Euchilopsis F. Muell.; Euchlora
Eckl. & Zeyh.; *Euchresta Benn.; Eutaxia R.Br. ex W.T.Aiton; Eversmannia Bunge; Exostyles Schott; Eysenhardtia Kunth; *Ezoloba B.-E.
van Wyk & Boatwr.; Fairchildia Britton & Rose; Fiebrigiella Harms; Fissicalyx Benth.; Flemingia Roxb. ex W.T.Aiton; Fordia Hemsl.; Galactia
P.Browne; Galega L.; Gastrolobium R.Br. ex W.T.Aiton; Geissaspis Wight & Arn.; Genista L.; Genistidium I.M.Johnston; Geoffroea Jacq.;
Gliricidia Kunth; Glycine Willd.; Glycyrrhiza L.; Gompholobium Sm.; *Gonocytisus Spach; Goodia Salisb.; Grazielodendron H.C.Lima;
*Greuteria Amirahm. & Kaz.Osaloo; Gueldenstaedtia Fisch.; Guianodendron Schutz Rodrigues & A.M.G.Azevedo; Halimodendron Fisch. ex
DC.; Hammatolobium Fenzl; Hanslia Schindl.; Haplormosia Harms; Hardenbergia Benth.; Harleyodendron R.S.Cowan; Harpalyce Moç. &
Sessé ex DC.; Haymondia A.N.Egan & B.Pan; Hebestigma Urb.; Hedysarum L.; Hegnera Schindl.; Helicotropis A.Delgado; *Herpyza C.Wright;
Hesperolaburnum Maire; Hesperothamnus Brandegee; Hippocrepis L.; Hoita Rydb.; Holocalyx Micheli; *Hosackia Douglas ex Lindl.; Hovea
R.Br. ex W.T.Aiton; Humularia P.A.Duvign.; Hylodesmum H.Ohashi & R.R.Mill.; Hymenocarpos Savi; Hymenolobium Benth.; Hypocalyptus
(Yakovlev) A.L.Schutte; Indigastrum Jaub. & Spach; Indigofera L.; Inocarpus J.R.Forst. & G.Forst.; Isotropis Benth.; Jacksonia R.Br. ex Sm.;
*Kebirita Kramina & D.D.Sokoloff; Kennedia Vent.; Kotschya Endl.; Kummerowia Schindl.; Kunstleria Prain; Lablab Adans.; Laburnum Fabr.;
Lackeya Fortunato, L.P.Queiroz & G.P.Lewis; Ladeania A.N.Egan & Reveal; Lamprolobium Benth.; Lathyrus L.; Latrobea Meisn.; *Lebeckia
Thunb.; Lecointea Ducke; *Lembotropis Griseb.; Lennea Klotzsch; Lens Mill.; Leobordea Delile; Leptoderris Dunn; *Leptodesmia (Benth.)
Benth.; Leptolobium Vogel; Leptosema Benth.; Leptospron (Benth. & Hook.f.) A.Delgado; Lespedeza Michx.; Lessertia DC.; Leucomphalos
Benth. ex Planch.; Limadendron Meireles & A.M.G.Azevedo; Liparia L.; *Listia E.Mey.; Lonchocarpus Kunth; Lotononis (DC.) Eckl. & Zeyh.;
Lotus L.; Luetzelburgia Harms; Lupinus L.; Luzonia Elmer; Maackia Rupr. & Maxim.; Machaerium Pers.; Macropsychanthus Harms; Macroptilium (Benth.) Urb.; Macrotyloma (Wight & Arn.) Verdc.; Maraniona C.E.Hughes, G.P.Lewis, Daza & Reynel ; Marina Liebm.; Mastersia
Benth.; Mecopus Benn.; Medicago L.; *Meizotropis Voigt; Melilotus Mill.; Melliniella Harms; Melolobium Eckl. & Zeyh.; Microcharis Benth.;
Mildbraediodendron Harms; Millettia Wight & Arn.; Mirbelia Sm.; Monarthrocarpus Merr.; Monopteryx Spruce ex Benth.; *Montigena
Heenan; Mucuna Adans.; Muellera L.f.; Muelleranthus Hutch.; Mundulea (DC.) Benth.; Myrocarpus Allemão; Myrospermum Jacq.; Myroxylon
L.f.; Mysanthus G.P.Lewis & A.Delgado; *Neocollettia Hemsl.; *Neoharmsia R.Vig.; Neonotonia J.A.Lackey; Neorautanenia Schinz; Neorudolphia Britton; Nephrodesmus Schindl.; Nesphostylis Verdc.; Neustanthus Benth.; Nissolia Jacq.; Nogra Merr.; Oberholzeria Swanepoel,
M.M.le Roux, M.F.Wojc. & A.E.van Wyk; Ohwia H.Ohashi; Olneya A.Gray; Onobrychis Mill.; Ononis L.; Ophrestia H.M.L.Forbes; Orbexilum
Raf.; *Oreophysa (Bunge ex Boiss.) Bornm.; Ormocarpopsis R.Vig.; Ormocarpum P.Beauv.; Ormosia Jacks.; Ornithopus L.; Orphanodendron
Barneby & J.W.Grimes; Oryxis A.Delgado & G.P.Lewis; *Ostryocarpus Hook. f.; Otholobium C.H.Stirt.; Otoptera DC.; *Ototropis Nees;
*Ottleya D.D.Sokoloff; *Ougeinia Benth.; Oxylobium Andrews; Oxyrhynchus Brandegee; Oxytropis DC.; Pachyrhizus Rich. ex DC.; Panurea
Spruce ex Benth.; Paracalyx Ali; *Paragoodia I.Thomps.; Paramachaerium Ducke; Paratephrosia Domin; Parochetus Buch.-Ham. ex D.Don;
Parryella Torr. & A.Gray ex A.Gray; Pearsonia Dummer; Pediomelum Rydb.; Periandra Mart. ex Benth.; Pericopsis Thwaites; Petaladenium
Ducke; Peteria A.Gray; Petteria C.Presl; Phaseolus L.; Philenoptera Fenzl ex A.Rich.; *Phylacium Benn.; Phyllodium Desv.; Phyllolobium
Fisch.; Phyllota (DC.) Benth.; Phylloxylon Baill.; Physostigma Balf.; Pickeringia Nutt. ex Torr. & A.Gray; Pictetia DC.; Piptanthus Sweet;
Piscidia L.; Pisum L.; Plagiocarpus Benth.; Platycelyphium Harms; Platycyamus Benth.; Platylobium Sm.; Platymiscium Vogel; Platypodium
Vogel; Platysepalum Welw. ex Baker; Podalyria Willd.; Podlechiella Maassoumi & Kaz.Osaloo; *Podocytisus Boiss. & Heldr.; Podolobium
R.Br. ex W.T.Aiton; *Podolotus Royle; Poecilanthe Benth.; Poiretia Vent.; *Poissonia Baill.; Poitea Vent.; Polhillia C.H.Stirt.; Pongamiopsis
R.Vig.; Pseudarthria Wight & Arn.; Pseudeminia Verdc.; Pseudoeriosema Hauman; *Pseudolotus Rech.f.; Pseudovigna (Harms) Verdc.;
Psophocarpus Neck. ex DC.; Psoralea L.; *Psoralidium Rydb.; Psorothamnus Rydb.; Pterocarpus Jacq.; Pterodon Vogel; Ptycholobium Harms;
*Ptychosema Benth.; Pueraria DC.; Pultenaea Sm.; Pycnospora R.Br. ex Wight & Arn.; *Pyranthus Du Puy & Labat; Rafnia Thunb.; Ramirezella Rose; Ramorinoa Speg.; Requienia DC.; Retama Raf.; Rhodopis Urb.; Rhynchosia Lour.; *Rhynchotropis Harms; Riedeliella Harms;
Robinia L.; *Robynsiophyton R.Wilczek; *Rothia Pers.; Rupertia J.W.Grimes; *Sakoanala R.Vig.; *Salweenia Baker f.; *Sarcodum Lour.;
Sartoria Boiss. & Heldr.; Schefflerodendron Harms; Scorpiurus L.; Securigera DC.; Sellocharis Taub.; Sesbania Adans.; Shuteria Wight &
Arn.; Sigmoidotropis (Piper) A.Delgado; Sinodolichos Verdc.; Smirnowia Bunge; Smithia Aiton; Soemmeringia Mart.; Solori Adans.; Sophora
L.; Spartium L.; Spathionema Taub.; Spatholobus Hassk.; Sphaerolobium Sm.; Sphaerophysa DC.; Sphenostylis E.Mey.; Sphinctospermum
Rose; Spirotropis Tul.; Spongiocarpella Yakovlev & N.Ulziykh.; Staminodianthus D.B.O.S.Cardoso, H.C.Lima & L.P.Queiroz; Stauracanthus
Link; Steinbachiella Harms; Stirtonanthus B.-E.van Wyk & A.L.Schutte; *Stonesiella Crisp & P.H.Weston; *Streblorrhiza Endl.; Strongylodon
Vogel; Strophostyles Elliott; Stylosanthes Sw.; Styphnolobium Schott; Sulla Medik.; Sutherlandia R.Br. ex W.T.Aiton; Swainsona Salisb.;
Swartzia Schreb.; Sweetia Spreng.; *Sylvichadsia Du Puy & Labat; *Syrmatium Vogel; Tabaroa L.P.Queiroz, G.P.Lewis & M.F.Wojc.; Tadehagi
H.Ohashi; Taralea Aubl.; Taverniera DC.; Templetonia R.Br. ex W.T.Aiton; Tephrosia Pers.; *Teramnus P.Browne; Tetragonolobus Scop.;
Teyleria Backer; Thermopsis R.Br. ex W.T.Aiton; Thinicola J.H.Ross; Tibetia (Ali) H.P.Tsui; Tipuana (Benth.) Benth.; Toxicopueraria A.N.Egan
& B.Pan; Trifidacanthus Merr.; Trifolium L.; Trigonella L.; Tripodion Medik.; Trischidium Tul.; Uleanthus Harms; Ulex L.; Uraria Desv.;
Uribea Dugand & Romero; Urodon Turcz.; *Vandasina Rauschert; Vatairea Aubl.; Vataireopsis Ducke; Vatovaea Chiov.; Vavilovia Al., Fed.;
*Verdesmum H.Ohashi & K.Ohashi; Vicia L.; Vigna Savi; Viminaria Sm.; Virgilia Poir.; *Vuralia Uysal & Ertuğrul; Wajira Thulin; Weberbauerella Ulbr.; *Wiborgia Thunb.; *Wiborgiella Boatwr. & B.-E.van Wyk; Wisteria Nutt.; Xanthocercis Baill.; *Xeroderris Roberty; Xiphotheca
Eckl. & Zeyh.; Zollernia Wied.-Neuw. & Nees; Zornia J.F.Gmel.; Zygocarpum Thulin & Lavin
Recent synonyms: Barnebyella Podlech = Astragalus L.; Bergeronia Micheli = Muellera L.f.; Calia Terán & Berland. = Dermatophyllum Scheele;
Etaballia Benth. = Pterocarpus Jacq.; Margaritolobium Harms = Muellera L.f.; Ophiocarpus (Bunge) Ikonn. = Astragalus L.; Paraderris (Miq.)
R.Geesink = Derris Lour.; Peltiera Labat & Du Puy = Ormocarpopsis R.Vig.; Spartidium Pomel = Calobota Eckl. & Zeyh.
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volume by Candolle (1825), Caesalpinioideae was chosen as
the preferred name for the MCC clade. Because of the broader
concept associated with Caesalpinioideae, it corresponds more
closely to the more inclusive MCC clade. Furthermore, this
leaves open the option in future classifications of naming the
morphologically distinct mimosoid clade at the tribal level and /
or under the International Code of Phylogenetic Nomenclature
(ICPN) (Cantino & de Queiroz, 2010).
In our new classification, three subfamily names are new
at this rank. We ascribe these names to the collective known
as the “Legume Phylogeny Working Group”. This uncommon
practice in botanical nomenclature does not prevent valid publication of the names under the botanical code as stipulated in
Chapter VI, Section 1 (Author Citations). Although we could
have adopted a modification of Recommendation 46C.2, which
suggests citing the first author followed by “et al.” (and at first
appearance of that authority, listing all 97 authors), we considered that ascribing authorship to the Legume Phylogeny
Working Group is more straightforward, more clearly gives
due credit to the legume systematics community and reflects
much better the collaborative approach used to arrive at this
new classification. At a time when systematics papers may
have increasing numbers of authors, for example, as genomic
datasets become routine, we feel that a desire for authorship
ascribed to research groups and communities rather than individuals will become more commonplace.
INTEGRATING TRIBAL AND CLADE-BASED
CLASSIFICATIONS
In addition to the need for a new Linnaean-based subfamily classification, there are important questions about the
best approach to naming clades within subfamilies. New phylogenies of many legume groups have unequivocally demonstrated the inadequacies of the tribal classifications of Polhill
& Raven (1981), Polhill (1994), and Lewis & al. (2005) because
of the non-monophyly of most of the traditionally recognised
tribes (LPWG, 2013a). In addition, questions remain about the
monophyly and placement of several genera, with considerable ongoing uncertainty surrounding generic delimitation and
relationships (LPWG, 2013a; Lewis & al., 2013). However,
numerous phylogenetic studies are ongoing and revised tribal
classifications of subfamilies will be forthcoming in the near
future. The emergence of clade-based phylogenetic classification systems provides an additional option to facilitate rankfree naming of robustly supported legume clades under the
draft ICPN. Such clade-based classifications can be easily
integrated with traditional Linnaean rank-based classification
to name additional clades coinciding with the evolution of key
biological traits that are hypothesised as synapomorphies. For
example, several important legume clades corresponding to
biologically important apomorphies (sometimes in the form
of deep homologies), including nodulation, bipinnate leaves
(here corresponding to the redefined Caesalpinioideae), extrafloral petiolar or leaf rachis nectaries, pollen in tetrads /
polyads, and valvate petal aestivation (mimosoid clade) could
66
be named in this way, as pursued by Wojciechowksi (2013)
for Papilionoideae using many of the recommendations of the
ICPN. We believe this approach, integrating Linnaean ranks
alongside clade-based ICPN classification, will greatly enhance
the biological meaning and utility of future classifications with
significant benefits for effective communication across a wide
spectrum of biological disciplines.
A new classification is clearly needed for the recircumscribed subfamily Caesalpinioideae, which has been the most
difficult and controversial to delimit in the new subfamilial
classification because of the inclusion of the formerly recognised and morphologically distinctive subfamily Mimosoideae.
Because relationships amongst major groups within the recircumscribed Caesalpinioideae remain poorly resolved (Figs.
1, 2 & S1), we refrain from establishing a new tribal and / or
clade-based classification for this subfamily here. Although
most mimosoids are morphologically distinct (Fig. 6), the
morphological distinctions between some members of the mimosoid clade and subtending caesalpinioid lineages are not
always clearcut. For example, Dinizia Ducke, once considered
to be in Mimosoideae, is placed outside the mimosoid clade in
molecular phylogenetic analyses (Luckow & al., 2003; Bruneau
& al., 2008), and Chidlowia Hoyle (Fig. 6A), which has always
been considered a caesalpinioid legume (Lewis & al., 2005), is
placed within the mimosoid clade in recent molecular phylogenetic analyses (Manzanilla & Bruneau, 2012; E.J.M. Koenen
& al., in prep.). For these reasons, we refrain from formally
naming this clade until relationships amongst lineages within
Caesalpinioideae can be better resolved, and refer to the former
subfamily Mimosoideae DC. simply as the “mimosoid clade”
for the time being.
In Cercidoideae and Dialioideae, both of which have relatively few genera (Lewis & Forest, 2005; Sinou & al., 2009;
E. Zimmerman, unpub. data), infra-subfamilial classifications
may not be needed, and Duparquetioideae is monospecific. In
Detarioideae, phylogenetic relationships amongst basal lineages have been too poorly resolved until now to permit their
classification (Bruneau & al., 2001, 2008; Fougère-Danezan
& al., 2007), but ongoing studies are leading to better resolution with the possibility for recognising clades as tribes
and / or formally named clades (M. de la Estrella & al., unpub.
data). Similarly, ongoing studies in Papilionoideae and in the
recircumscribed Caesalpinioideae should help resolve key relationships, with the ultimate outcome that names of strongly
supported and biologically meaningful clades will be proposed
in forthcoming publications.
REFERENCE PHYLOGENY
The classification proposed here uses as its framework
the most comprehensively sampled phylogenetic analysis of
legumes to date (Figs. 1, S1; Table S1; Methods described in
Appendix 1). This new phylogeny is based on plastid matK
gene sequences because this gene region is the most widely
sequenced across the legumes (cf. LPWG, 2013a) and it is
sufficiently variable to resolve generic membership of many
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strongly supported higher-level clades as demonstrated by
a large number of studies such as those referenced herein.
Although this analysis is based on a single plastid locus, the
topology observed and the groups that are supported have been
consistently resolved in numerous previous phylogenetic analyses of the entire family or of particular clades within the family
using diverse plastid (trnL-F, trnD-T, rbcL, rps16, rpl16) and
nuclear loci (e.g., rDNA ITS, SucS) (see LPWG, 2013a and
references therein). In recent analyses of all 81 plastid genes
(Fig. 2) and of a large nuclear gene dataset derived from transcriptome sequences (E.J.M. Koenen & al., in prep.), all five of
the non-monospecific subfamilies are strongly supported, and
the relationships amongst them do not conflict with the matK
analyses (see below), although the nuclear gene dataset does
not include Duparquetioideae.
The analysis presented here includes 3696 legume species
(with an additional 48 infraspecific taxa) representing 698 of
the currently recognised 765 legume genera (Figs. 1 & S1;
Tables 2 & S1; Appendix 1). Subfamilies Cercidoideae and
Duparquetioideae are fully sampled at the generic level. In the
Detarioideae, five genera are not sampled, all of them monospecific, in Dialioideae two monospecific genera are missing, and
in Caesalpinioideae, two genera are not sampled (Table 2, missing genera identified with *). Papilionoideae are represented by
445 genera, with most of the missing 48 genera belonging to
the tribe Loteae and phaseoloid clades. The phylogenetic trees
and the underlying alignment and voucher data are available
to browse and download from the online Supplementary Data
(Table S1; Data Files A–F) and on Data Dryad (DOI: https://
doi.org/10.5061/dryad.61pd6).
Bayesian analyses (Fig. 1) and maximum likelihood (Fig.
S1) of the matK sequence data resolve the Leguminosae as
monophyletic with 1.0 posterior probability and 100% bootstrap support. Each of the five non-monospecific subfamilies
of Leguminosae is also supported with 1.0 posterior probability and 100% bootstrap support. Relationships amongst subfamilies Cercidoideae, Detarioideae, Duparquetioideae and
the clade that groups the remaining legumes (i.e., the other
three subfamilies) are unresolved, forming a basal polytomy
(Fig. 1). Dialioideae is resolved as sister to Caesalpinioideae +
Papilionoideae (1.0 posterior probabability, Fig. 1; 100%
bootstrap support, Fig. S1), which are sister to each other. In
the full plastid analyses of E.J.M. Koenen & al. (in prep.),
Duparquetioideae is robustly supported as sister to the
Dialioideae + Caesalpinioideae + Papilionoideae clade, but the
relationship of this clade to the Cercidoideae and Detarioideae
remains unresolved (Fig. 2). Many genera of Leguminosae are
supported as monophyletic in the matK analysis, with notable
exceptions for certain large genera that are the focus of ongoing taxonomic and phylogenetic studies (e.g., Bauhinia s.l. in
Cercidoideae, several genera of Detarioideae, of the mimosoid clade, and of tribe Millettieae in Papilionoideae). In the
mimosoid clade, and in other parts of Caesalpinioideae and
Detarioideae, genera are often not supported as monophyletic,
and generic-level relationships are often poorly resolved. This
can likely be attributed in part to striking substitution rate
heterogeneity in plastid genes, and hence variable phylogenetic
resolution across legumes, as previously noted by Lavin & al.
(2005) and Dugas & al. (2015) (see also Figs. 1 & 2).
Several recent large-scale angiosperm / rosid phylogenetic
analyses (Zanne & al., 2014; Li & al., 2016; Sun & al., 2016)
included thousands of legume nuclear and plastid and, in
some cases, mitochondrial sequences. These analyses contain
many taxa that were mis-identified or labelled using outdated
taxon names, or are based on apparent sequence contaminants
that have been deposited in GenBank without being properly
checked and annotated. These inaccuracies, compounded by
large amounts of missing data (e.g., 80% in Zanne & al., 2014),
together interact to cause unpredictable and chaotic problems in
phylogenetic analyses, a phenomenon highlighted several years
ago by McMahon & Sanderson (2006) in their supermatrix
phylogenetic analysis of papilionoid legumes. Unfortunately,
such potentially flawed topologies have been used as the basis
for several recent large-scale evolutionary studies focused, for
example, on key characteristics of legumes such as the origins of nodulation and nitrogen fixation (e.g., Werner & al.,
2014, 2015; Li & al., 2016). A cursory examination of many of
these large-scale phylogenies has revealed a number of unusual
and demonstrably inaccurate relationships. Using such badly
flawed phylogenies can obviously lead to weak or even erroneous conclusions regarding the evolution of particular traits
(cf. Doyle, 2016). In contrast, the phylogeny presented here
uses a fully curated set of sequences that are vouchered and
taxonomically validated by the legume systematics community.
The phylogenetics of legumes, like that of any major clade,
is of course a work in progress. New densely sampled phylogenies at the species, generic and higher levels based on full
plastome sequences, as well as transcriptomes and hundreds
of nuclear loci are ongoing, and will in due course supersede
the phylogeny presented here. Regardless, the taxonomically
validated tree presented here can be used for downstream analyses that require an accurate and densely sampled phylogenetic
framework of the Leguminosae.
TAXONOMY
Based on the phylogenetic structure of the family
Leguminosae presented here, we recognise six subfamilies.
We provide a key, taxonomic descriptions for each of the
subfamilies, and illustrate the diversity of flowers and fruits
across these subfamilies (Figs. 3–11). Comparative morphology, chemistry and chromosome numbers of the six subfamilies
(Table 1) and a full list of genera by subfamily, noting recent
synonyms (Table 2) are presented. Technical terms are defined
and illustrated in Appendix 2 and Figs. 12 & 13.
Key to the subfamilies of Leguminosae
1.
1.
Petals with marginal glandular structures; flowers with
4 stamens, anthers fused in a synandrium with poricidal
dehiscence; leaves once pinnate; endemic to Central and
West Africa .............................. Duparquetioideae
Petals not glandular (except in the Amazonian, papilionoid
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2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
genus Petaladenium); flowers with 4 stamens uncommon
(but then anthers never fused in a synandrium); anther dehiscence longitudinal (except poricidal in some genera of
Caesalpinioideae); leaves various; widely distributed ... 2
Flowers mostly papilionate (“pea-flowered”) and bilaterally symmetrical, less commonly radially symmetrical;
median (standard) petal outermost, enclosing the wing and
keel petals (especially in bud) or the wing and keel petals
lacking; sepals united, at least at the base, into a calyx
tube or completely enclosing the floral bud; seeds with a
complex hilar valve, pleurogram absent; embryo radicle
usually curved .............................. Papilionoideae
Flowers not papilionate (if rarely appearing papilionate then the median petal innermost), either bilaterally
or radially symmetrical, median (standard) petal innermost, or petals valvate (in the mimosoid clade of the
Caesalpinioideae); sepals free or fused; seeds lacking
complex hilar valve, with or without a pleurogram; embryo radicle usually straight ................................ 3
Leaves bipinnate; seeds commonly with an open or closed
pleurogram on each side ................ Caesalpinioideae
Leaves never bipinnate; seeds without an open or closed
pleurogram on either side ................................... 4
Leaves unifoliolate, bilobed or entire, or compound and
bifoliolate; seed hilum circular or crescent-shaped .......
................................................... Cercidoideae
Leaves various; if simple or bifoliolate, then the seed hilum
not crescent-shaped, and rarely circular .................. 5
Extra-floral nectaries and other glandular structures (when
present) on the lower surface or margin of leaflets; stipules usually intrapetiolar (free or united), rarely lateral
.................................................... Detarioideae
Extra-floral nectaries absent or present on the petiole or
on the leaf rachis; stipules lateral and free or absent .... 6
Inflorescences highly branched and thyrsoid or racemes
with distichous anthotaxy; leaves mostly imparipinnate
with alternate leaflets (rarely paripinnate with oppostite
leaflets in Eligmocarpus and Poeppigia), extra-floral nectaries on the petiole or leaf rachis absent .... Dialioideae
Inflorescences mostly racemes with spiral anthotaxy,
commonly compounded into branched panicles or contracted in spikes or fascicles; leaves mostly paripinnate
with opposite leaflets, rarely bifoliolate or with alternate
leaflets; extra-floral nectaries (when present) on the petiole or on the leaf rachis between the leaflet pairs .......
.............................................. Caesalpinioideae
Descriptions of the six subfamilies
A short description is presented for each subfamily, highlighting (in bold) the diagnostic features of each.
Subfam. Cercidoideae Legume Phylogeny Working Group,
stat. nov. ≡ Cercideae Bronn, Form. Pl. Legumin.: 134,
131. 1822 (“Cerceae”) – Type: Cercis L.
Trees, shrubs or tendriled lianas (Figs. 3A–F), mostly
unarmed but frequently with prickles or infrastipular spines,
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branches rarely modified into flattened cladodes (Brenierea
Humbert); specialised extrafloral nectaries stipular when present (Bauhinia L.), never on petiole and leaf rachis. Stipules in
lateral position, free. Leaves uni- or bifoliolate (bipinnate,
pinnate, palmate and trifoliolate leaves totally lacking),
pulvinate, leaflet blade (when unifoliolate) entire or bilobed
with a small mucro at the apex or between the lobes, exstipellate. Inflorescence racemose or pseudoracemose; bracteoles
minute or large. Flowers bisexual, rarely unisexual (plants
polygamous or dioecious), slightly to strongly bilaterally symmetrical, sometimes papilionate (Cercis), hypanthium greatly
elongated to almost absent; sepals united in a spathaceous
or 2–5-lobed calyx or free; petals free, 5, rarely 2, 6 (some
Bauhinia) or absent (Brenierea), imbricate, the adaxial petal
innermost and frequently differentiated; stamens usually 10
(sometimes fewer) in two whorls of alternate length, the filaments partly connate or free, anthers mostly uniform and
dorsifixed, opening by a longitudinal slit or central pore in
each theca, reduced stamens or staminodes sometimes present; pollen 3-colporate, 3–6-colpate, 3-porate, 3-pororate,
3–4-colporoidate or inaperturate monads, rarely in tetrads
(only in some Bauhinia); gynoecium 1-carpellate, stipe of
ovary free or adnate to abaxial wall of the hypanthium, ovary
1–many-ovulate. Fruits dehiscent (often explosively with twisting valves) or indehiscent and then generally samaroid. Seeds
with apical crescent-shaped hilum, rarely circular (Cercis),
lens inconspicuous, lacking pleurograms, pseudopleurograms,
or wing or aril (in Brenierea two funicular aril-like lobes adnate to the testa leaving a short crescent-shaped scar or a long
scar running nearly around the seed circumference); embryo
straight, very rarely curved (Barklya F.Muell.). Vestured pits
lacking in secondary xylem; silica bodies absent; septate fibres
and storeyed rays sometimes present. Root nodules absent.
2n = 14, 24, 26, 28 (42, 56). Coumarins and cyanogenic glucosides reported; non-protein amino acids common (5-hydroxy-L-tryptophan only reported in this subfamily, Griffonia
Baill., Brenierea).
Currently 12 genera and ca. 335 species, mainly tropical,
Cercis in the warm temperate Northern Hemisphere.
Clade-based definition (included taxa): The most inclusive
crown clade containing Cercis canadensis L. and Bauhinia divaricata L. but not Poeppigia procera C.Presl, Duparquetia orchidacea Baill., or Bobgunnia fistuloides (Harms) J.H.Kirkbr.
& Wiersema.
Subfam. Detarioideae Burmeist., Handb. Naturgesch.: 319.
1837 (“Detarieae”) – Type: Detarium Juss.
Unarmed trees, sometimes shrubs, rarely suffruticose
(Cryptosepalum Benth.) (Fig. 4); specialised extrafloral nectaries often present abaxially, rarely on the margins of leaflets
or on leaf rachis, and never on the petiole. Stipules in intrapetiolar position (i.e., somewhere between the petiole and
the axillary bud) and then free, valvate and connected by
chaffy hairs, or fused, either partly (only at base) or entirely, rarely lateral and free. Leaves paripinnate (ending in a
pair of leaflets or, if leaflets alternate and appearing imparipinnate, the terminal leaflet exceeded by a more or less caducous
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rachis-extension) with 1 (bifoliolate) to numerous pairs of
leaflets, or rarely unifoliolate (Paloue Aubl., Paloveopsis
R.S.Cowan, Zenkerella Taub., some Cryptosepalum, Didelotia
Baill. and Guibourtia Benn.), bipinnate leaves totally lacking,
leaves pulvinate, leaflets opposite or alternate, exstipellate,
translucent glands sometimes present. Inflorescence a raceme
or panicle; bracteoles small to large, frequently petaloid, valvate or imbricate, free or partially fused or partly fused with
the hypanthium, partially or completely enclosing the bud.
Flowers bisexual or with both bisexual and male flowers radially or slightly to strongly bilaterally symmetrical (but never
papilionate), hypanthium elongated to almost absent; sepals
commonly 5 or 4 (two adaxial sepals often fused), rarely some
or all absent or more (–7); petals free, 0–5(–7), when present
imbricate, the adaxial petal generally innermost (outermost in
some flowers of Hymenaea L. and allies), all equal or the adaxial large and either the other 4 or only the abaxial ones smaller
to rudimentary; stamens 2–numerous but usually 10, the filaments partly connate or free, staminodes occasionally present;
anthers dorsifixed or basifixed; pollen mostly 3-colporate monads with a vast array of sculptures; gynoecium 1-carpellate,
1–many ovulate, stipe of ovary free or adnate to the wall of the
hypanthium. Fruits mostly woody, dehiscent pods, sometimes
indehiscent and woody or thin-valved, samaroid (Brandzeia
Baill., Barnebydendron J.H.Kirkbr., Gossweilerodendron
Harms, Hardwickia Roxb., Neoapaloxylon Rauschert), rarely
filled with pulpy mesocarp (Tamarindus L.) or endocarp
(Hymenaea). Seeds often overgrown, sometimes hard and
then occasionally with pseudopleurograms (Lysidice Hance,
Paramacrolobium J.Léonard, Peltogyne Vogel, Tamarindus),
occasionally arillate; embryo straight. Vestured pits present
in secondary xylem; axial (resin) canals sometimes present;
silica bodies rarely present (Hymenostegia Harms, Loesenera
Harms); septate fibres and storeyed rays sometimes present.
Root nodules absent. 2n mostly 24 but occasionally 16, 20, 22,
36, 68. Coumarins reported; frequently with terpenes (resins)
and non-protein amino acids.
Currently 84 genera and ca. 760 species, almost exclusively
tropical, Schotia Jacq. in sub-tropical South Africa.
Clade-based definition (included taxa): The most inclusive
crown clade containing Goniorrhachis marginata Taub. and
Aphanocalyx cynometroides Oliv., but not Cercis canadensis,
Duparquetia orchidacea or Bobgunnia fistuloides.
Subfam. Duparquetioideae Legume Phylogeny Working
Group, stat. nov. ≡ Duparquetiinae H.S.Irwin & Barneby
in Polhill & Raven, Adv. Legume Syst. 1: 102. 1981 – Type:
Duparquetia Baill.
Unarmed, scrambling liana (Fig. 3G), often climbing to the
forest canopy; specialised extrafloral nectaries lacking on petiole and leaf rachis. Stipules in lateral position, free, narrowly
triangular. Leaves imparipinnate, pulvinate, leaflets opposite,
exstipellate. Inflorescence a terminal, erect, 10–30-flowered
raceme; bracteoles 2, small. Flowers bisexual, strongly bilaterally symmetrical, hypanthium lacking; sepals 4, unequal, the
abaxial and adaxial sepals cucullate, sepaloid, the lateral
sepals petaloid; petals 5, free, dimorphic, the adaxial and the
two lateral petals ovate, two abaxial petals strap-like, oblong,
all 5 petals with stalked gland-like extrusions along their
margins, imbricate, the adaxial petal outermost; stamens 4,
the anthers basifixed, oblong, with pointed appendages, the
thecae dehisce by a short, apical, poricidal slit, the anthers
postgenitally fused into a curving synandrium, the appendages remain free; pollen in monads, asymmetrical, one
equatorial-encircling ectoaperture with two equatorial endoapertures; gynoecium 1-carpellate, stipitate, 2–5-ovuled,
with four ridges running along the length of the ovary. Fruit
an oblong four-angled, woody pod, dehiscent, valves spirally
coiled. Seeds 2–5 per fruit, oblong to ovoid, the testa thick; embryo straight. Vestured pits lacking in secondary xylem; silica
bodies, septate fibres and storeyed rays absent. Root nodules
absent. Chromosome number unknown.
Monospecific: Duparquetia orchidacea Baill. Distributed
in humid tropical forests of West and Central Africa.
Subfam. Dialioideae Legume Phylogeny Working Group, stat.
nov. ≡ Dialiinae H.S.Irwin & Barneby in Polhill & Raven,
Adv. Legume Syst. 1: 100. 1981 – Type: Dialium L.
Unarmed trees or shrubs, rarely suffruticose (Labichea
Gaudich. ex DC., Petalostylis R.Br.) (Fig. 3H–L); specialised
extrafloral nectaries lacking on petiole and leaf rachis and
on leaflet surface. Stipules in lateral position, free or absent.
Leaves imparipinnate, rarely paripinnate (Eligmocarpus
Capuron, Poeppigia C.Presl), 1-foliolate (Baudouinia Baill.,
Labichea, Mendoravia Capuron, Uittienia Steenis) or palmately compound (Labichea), leaflets alternate, rarely opposite (Eligmocarpus, Poeppigia), exstipellate. Inflorescences
highly branched, thyrsoid, less commonly racemes with distichous anthotaxy (Labichea, Petalostylis), borne in both terminal and axillary positions, or reduced to one axillary flower
(Petalostylis); bracteoles small or absent. Flowers bisexual
(polygamous in Apuleia Mart.), radially or slightly to strongly
bilaterally symmetrical, hypanthium rarely present, receptacle may be broad and flattened, bearing nectary-like bodies; sepals commonly 5, reduced to 4 (Labichea, Storckiella
Seem.) or 3 (Apuleia, Dialium), rarely 6 (Mendoravia), free,
equal to sub-equal; petals 5 or fewer (0, 1, 3, 4), rarely 6 (petal
number often equivalent to sepal number), free, equal to subequal, imbricate, the adaxial petal innermost; fertile stamens
5 or fewer, rarely 6–10 (some Dialium spp., Poeppigia), usually only antesepalous whorl present, free, uniform, rarely
dimorphic (Eligmocarpus), anthers basifixed, rarely dorsifixed (Poeppigia), dehiscing via longitudinal slits, often
reduced to a short apical, poricidal slit, staminodes present or absent; pollen in tricolporate monads with punctate
or finely reticulate, rarely striate (some Dialium) sculpture
patterns; gynoecium 1-carpellate (sometimes bicarpellate
in scattered flowers of Dialium), ovary stipitate or sessile,
ovules frequently 2 (1–many). Fruits commonly indehiscent
drupaceous or samaroid, rarely dehiscent (Eligmocarpus,
Labichea, Mendoravia, Petalostylis) or the drupaceous fruit
with indurating endocarp breaking up in one seeded segments (Baudouinia). Seeds 1–2, rarely more; embryo straight.
Vestured pits absent in the secondary xylem, rarely present
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(Poeppigia, Mendoravia); silica bodies sometimes present
(Apuleia, Dialium, Dicorynia Benth., Distemonanthus Benth.);
septate fibres rarely present (Apuleia, Distemonanthus,
Poeppigia); storeyed rays often present. Root nodules absent.
2n = 28 (most genera unsurveyed).
Currently 17 genera and ca. 85 species. Widespread
throughout the tropics, with taxa occurring in South, Central
and North America, Africa, Madagascar, South and Southeast
Asia, south China, Australia, New Guinea and some Pacific
islands.
Clade-based definition (included taxa): The most inclusive
crown clade containing Poeppigia procera and Dialium guianense (Aubl.) Sandwith, but not Cercis canadensis, Duparquetia
orchidacea, or Bobgunnia fistuloides.
Subfam. Caesalpinioideae DC., Prodr. 2: 473. 1825 – Type:
Caesalpinia L.
= Mimosoideae DC., Prodr. 2: 424. 1825 – Type: Mimosa L.
= Cassioideae Burmeist., Handb. Naturgesch.: 319. 1837
(“Cassieae”) – Type: Cassia L., nom. cons.
Trees, shrubs, lianas, suffruticose or functionally herbaceous, occasionally aquatic (Figs. 5 & 6), either unarmed or
commonly armed with prickles or nodal or infranodal spines;
specialised extrafloral nectaries often present on the petiole and / or on the primary and secondary rachises, usually between pinnae or leaflet pairs, more rarely stipular
or bracteal (Senna Mill., Macrosamanea Britton & Rose ex
Britton & Killip and some Archidendron F.Muell.). Stipules
in lateral position and free or absent. Leaves usually pulvinate, commonly bipinnate, otherwise pinnate (sometimes
both types on the same plant in Arcoa Urb., Cenostigma Tul.,
Gleditsia L., Stuhlmannia Taub., rarely in Ceratonia L. and
Moldenhawera Schrad.) and then mostly paripinnate, rarely
imparipinnate, less often bifoliolate, modified into phyllodes
or lacking, arrangement of the pinnae and leaflets mostly opposite, rarely alternate; stipels rare and not to be confused with
the more commonly present paraphyllidia. Inflorescences
globose, spicate, paniculate, racemose or in fascicles; bracteoles commonly absent or small. Flowers usually bisexual,
rarely unisexual (Ceratonia, Gleditsia and Gymnocladus Lam.,
species dioecious or monoecious), or bisexual flowers combined with unisexual and / or sterile flowers in heteromorphic
inflorescences (mimosoid clade), radially, less frequently bilaterally symmetrical or asymmetric, hypanthium lacking or
cupular, rarely tubular; sepals (3–)5(–6), free or fused; petals
(3–)5(–6), free or fused (the sepal or petal or both whorls sometimes lacking), aestivation valvate (mimosoid clade) or imbricate and then the adaxial petal innermost; stamens commonly
diplostemonous or haplostemonous, sometimes reduced to 3,
4 or 5 (in some Mimosa spp.), frequently many (100+ in some
mimosoids), free or fused, sometimes heteromorphic, some
or all sometimes modified or staminodial, anthers basifixed
or dorsifixed, often with a stipitate or sessile apical gland,
dehiscing via longitudinal slits or apical or basal poricidal slits
or pores; pollen in tricolporate monads, or commonly in tetrads, bitetrads or polyads (most mimosoids); gynoecium unior rarely polycarpellate, 1–many-ovulate. Fruit a thin-valved,
70
1–many-seeded pod, dehiscent along one or both sutures, also
often a lomentum, a craspedium, or thick and woody and then
indehiscent or explosively dehiscent, often curved or spirally
coiled. Seeds usually with an open or closed pleurogram
on both faces, sometimes with a fleshy aril (Pithecellobium
Mart. and some Acacia Mill.) or sarcotesta (Inga Mill.), sometimes winged; hilum usually apical, lens usually inconspicuous;
embryo straight. Vestured pits present in secondary xylem;
silica bodies sometimes present (Tachigali Aubl., Diptychandra
Tul.); septate fibres and storeyed rays sometimes present. Root
nodules variably present and indeterminate (prevalent in
the mimosoid clade). 2n mostly 24, 26, 28, but also reported
2n = 14, 16, 52, 54, 56. Non-protein amino acids frequently reported, for example mimosine, albizine (mimosoids), djenkolic
acid, pipecolic acid and its derivatives; coumarins, cyanogenic
glucosides, phenylethylamines, tryptamines, and β-carboline
alkaloids also reported.
Caesalpinioideae in its emended circumscription contains
148 genera and ca. 4400 species. Pantropical, common in both
wet and dry regions, with a handful of species extending to
the temperate zone, less frequently frost-tolerant (Gleditsia,
Gymnocladus and some species of Desmanthus Willd. and
Senna).
This clade was referred to as the MCC clade (Doyle, 2011,
2012) or GCM-clade (Marazzi & al., 2012).
Clade-based definition (included taxa): The most inclusive
crown clade containing Arcoa gonavensis Urb. and Mimosa
pudica L., but not Bobgunnia fistuloides, Duparquetia orchidacea, or Poeppigia procera.
Subfam. Papilionoideae DC., Prodr. 2: 94. 1825 ≡ Faboideae
Rudd in Rhodora 70: 496. 1968 – Type: Faba Mill., (≡
Vicia L.).
= Lotoideae Burnett, Outlines Bot.: 643. 1835 (“Lotidae”) –
Type: Lotus L.
Mostly unarmed trees, shrubs, lianas, herbs, and twining
or tendriled vines (Figs. 7–9); specialised extrafloral nectaries
lacking on petiole and leaf rachis, occasionally stipular, stipellar or bracteal nectaries, or swollen and nectar-secreting peduncles, rarely on sepals (Erythrina L.). Stipules in lateral position (very rarely interpetiolar, in all species of Platymiscium
Vogel), free or absent. Leaves mostly pari- or imparipinnate
to palmately compound, also commonly uni- or trifoliolate,
rarely bi- or tetrafoliolate, never bipinnate (palmately-pinnate
in Rhynchosia ferulifolia Benth. ex Harv.), either pulvinate
or not, leaflets opposite or alternate, sometimes modified
into tendrils, rarely phyllodinous, stipels present or absent.
Inflorescence mostly racemose, pseudoracemose or paniculate,
less often cymose, spicate or capitate, axillary or terminal, or
flowers solitary; bracteoles usually present, rarely enlarged,
valvate, enveloping bud. Flowers bisexual, rarely unisexual,
usually bilaterally symmetrical, rarely asymmetrical, radially
symmetrical or nearly so, rarely cleistogamous flowers also
present; hypanthium present or absent; sepals (3–)5, united
at least at the base, sometimes the calyx entire and splitting
into irregular lobes or the calyx lobes dimorphic and some
petaloid; petals (0–)5(–6) and then imbricate, corolla mostly
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LPWG • Phylogeny and classification of the Leguminosae
papilionate, with the adaxial petal (= standard) outermost
and largest, usually overlapping lateral wing petals which in
turn overlap the abaxial keel petals or, in radially symmetrical
flowered species, corolla with 5 small or undifferentiated petals, less often only one (standard) petal is present or all petals
absent; stamens typically 10, rarely 9 or many, filaments most
commonly connate into a sheath or tube, or uppermost filament wholly or partly free, sometimes all filaments free, anthers uniform or dimorphic, basifixed or dorsifixed, dehiscing
longitudinally; pollen in monads, mostly 3-colporate, 3-colpate
or 3-porate; gynoecium 1-carpellate, very rarely 2-carpellate,
1–many-ovuled. Fruit a 1–many seeded pod, dehiscing along
one or both sutures, or indehiscent, or a loment, samara or
drupe. Seeds usually with a hard testa, rarely overgrown, sometimes with a fleshy aril or sarcotesta, a complex hilar valve,
elongate hilum and lens usually present, pleurogram absent;
embryo usually curved, rarely straight. Vestured pits present
in secondary xylem; silica bodies absent; septate fibres sometimes present; all elements (vessels, parenchyma, strands rays)
usually in storeyed structure. Root nodules generally present,
either indeterminate or determinate. 2n = more commonly
16, 18, 20, 22 but other numbers also reported (2n = 12, 14, 24,
26, 28, 30, 32, 38, 40, 48, 64, 84). Isoflavonoids, prenylated
flavonoids, indolizidine or quinolizidine alkaloids reported.
Non-protein amino acids widespread, some exclusively found
in the subfamily (e.g., canavanine).
Currently 503 genera and ca. 14,000 species, nearly
cosmopolitan.
Clade-based definition (included taxa): The most inclusive
crown clade containing Castanospermum australe A.Cunn.
ex Mudie and Vicia faba L., but not Erythrostemon gilliesii
(Hook.) Klotzsch., Gleditsia triacanthos L., or Dialium guianense. For ICPN classification of particular Papilionoideae
clades see Wojciechowski (2013).
The mimosoid clade
Although the mimosoid clade (Fig. 6) is not formally
named here, it is morphologically distinct and can be defined
as the most inclusive crown clade containing all Leguminosae
with radially symmetrical flowers having valvate petal aestivation, homologous to those found in Pentaclethra macrophylla
Benth. and Inga edulis Mart.
The mimosoid clade contains all genera previously assigned to subfamily Mimosoideae plus Chidlowia, previously
considered to be a member of the former Caesalpinioideae,
but now shown to belong to the mimosoid clade (Manzanilla
& Bruneau, 2012; E.J.M. Koenen & al., in prep.). This clade
of 3300+ species is morphologically highly distinctive with
radially symmetrical flowers with valvate aestivation of the
calyx and corolla (except in Parkia, which has partially imbricate calyx lobes). Typically, flowers are combined in spicate or
capitate inflorescences, often these are in turn combined into
compound inflorescences (e.g., a panicle of globose heads).
Pantropical, common in both wet and dry regions, with a
handful of species extending to the temperate zone, and less
frequently into frost-prone regions.
ACKNOWLEDGEMENTS
This project was made possible thanks to ongoing collaboration
with researchers worldwide who have collected samples of legumes
for DNA studies and who have shared material with members of the
legume systematics community for over three decades. In particular,
we thank the following people for their help with collection, preparation and curation of specimens, and / or with sequencing of the matK
locus: Alexandra Clark, Michelle Hart (Royal Botanic Gardens,
Edinburgh); Adilson M. Pintor, Marcelo T. Nascimento, Pablo Prieto
(Rio de Janeiro); Aécio A. Santos (Goiás, Tocantins); Antônio S.L. da
Silva, Camilo Barbosa, Catarina S. Carvalho, Leandro V. Ferreira,
Lisandra A. Teixeira, Nara Mota, Pedro L. Viana, Rafael Salomão
(Pará); Caio V. Vivas, José Lima Paixão, Tim Baker (Bahia); Claudio
Nicoletti, Geovane S. Siqueira (Espirito Santo); Eric Hattori, Fernanda
S. Freitas, Flávia Pezzini, Pedro Taucce (Minas Gerais); Marcella Baroni
(Mato Grosso do Sul); Flávia Costa (Manaus); Luzmilla Arroyo, Daniel
Villaroel, Alexander Germaine Parada (Bolivia); Aniceto Daza, Jose
Luis Marcelo-Peña, Reynaldo Linares Palomino, Carlos Reynel, Isau
Huamantupa (Peru); Tanja Schuster, Mansi Trivedi, Gabe Johnson,
William Cagle, Ailsa Holland, and Xin-Fen Gao (Smithsonian
Institution). We thank the ScienceCloud at the University of Zurich for
computational resources. Funding for this project was provided by the
Natural Sciences and Engineering Research Council of Canada, the U.K.
Natural Environment Research Council (grant NE/I028122/1), the Swiss
National Science Foundation (grant 31003A_13552), the Coordenação
de Aperfeiçoamento de Pessoal de Nível Superior (CAPES / Program
POS CSF # 1951/13-0), Conselho Nacional de Desenvolvimento
Científico e Tecnológico (Project Sisbiota 563084/2010-3 and Project
Casadinho/Procad # 5525892011-0), Fundação de Amparo à Pesquisa
do Estado da Bahia (FAPESB PES 0053/2011) and the Fundação de
Amparo à Pesquisa do Estado de São Paulo FAPESP; Conselho Nacional
de Desenvolvimento Científico e Tecnológico CNPq of Brazil, the
Smithsonian Institution, the U.S. National Science Foundation (grant
DEB-1352217), and the Environment Research and Technology
Development Fund (S9) of the Ministry of the Environment of Japan,
and Arizona State University. Anne Bruneau acknowledges the Royal
Botanic Gardens, Edinburgh, and the Department of Systematic and
Evolutionary Botany, University of Zurich for logistical support during
a sabbatical in 2015. Finally, we thank Jonathan Amith, Xander van der
Burgt, Emilio Constantino, David Du Puy, Flora do Acre, Felix Forest,
Paul Hoekstra, Mike Hopkins, Rosangela Melo, Justin Moat, Projecto
Flora Reserva Ducke INPA/DFID, Shijin Li, Alex Popovkin, James
Ratter, Wolfgang Stuppy, Liam Trethowan, Timothy Utteridge, and
André van Proosdij for contributing excellent legume images to Figs.
3–11. The authors thank Michael Pirie, Lars Chatrou, Jefferson Prado,
John McNeill, and two anonymous reviewers for advice on nomenclatural issues and comments on the manuscript.
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Appendix 1. Materials and Methods: A densely sampled phylogeny of the Leguminosae based on analyses of matK gene sequences.
Sampling. — Previously published and 637 newly generated matK gene sequences were obtained from multiple laboratories. Only fully vouchered
samples, authoritatively identified by taxonomic specialists are included, and
all sequences have been submitted to GenBank (Table S1). Most sequences
comprise the full matK coding sequence, but for a subset of species only
620–780 nucleotides of the central gene region, the “barcode” matK region
(from ca. 600 to 1450 in the aligned sequence matrix), were available. Our
objective was to include as many legume genera and species as possible,
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while at the same time ensuring sequence quality and taxonomic accuracy.
Multiple accessions per species were included in initial phylogenetic
analyses (all accessions listed in Table S1) in order to verify sequence accuracy
and try to eliminate problems of sequence contamination or specimen taxonomic identity. A total of 5560 legume sequences were verified and analysed.
Most sequences were also subjected to a BLAST search (http://blast.ncbi.nlm.
nih.gov/) to verify sequence accuracy. Subsequently a single sequence per
species (or infraspecific taxon) was chosen for the full phylogenetic analyses
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Appendix 1. Continued.
(specimens selected in the single-taxon analyses are indicated by * in Table
S1). Full-length gene sequences were preferentially selected. The aligned
matrix, accession list with voucher information and GenBank numbers, and
tree files (Bayesian and bootstrap majority-rule consensus trees, best-scoring
ML tree, 1000 Bayesian posterior trees and 1000 bootstrapped trees, all in
newick format) are available as Supplementary Data and in Data Dryad (DOI:
https://doi.org/10.5061/dryad.61pd6).
The final matrix includes 3842 sequences representing 3696 legume
species (identified with * in Table S1) and covering 698 of the 765 currently
recognised legume genera (Table 2). The sampling for subfamily Cercidoideae
includes 96 species representing all 12 genera; for Detarioideae, 327 species (plus 3 infraspecific taxa) are included from 79 of the 84 genera; for
Dialioideae, 19 species are included, representing 15 of the 17 genera; for
Caesalpinioideae, we include 937 species (plus 5 infraspecific taxa), representing 146 of the 148 genera; and for Papilionoideae, 2316 species are
included (plus 38 infraspecific taxa), representing 445 of the 503 genera.
This represents the most comprehensive generic and species sampling of
Leguminosae in a phylogenetic analysis of the family to date.
We also included 100 outgroup sequences, sampled across Eudicots,
including relatively dense sampling of the three other families of Fabales
(Table S1). Broad sampling of outgroup taxa was included to facilitate downstream analyses requiring branch lengths and wider interfamilial relationships. Sequences for outgroup taxa were obtained from vouchered GenBank
sequences (including published complete plastome sequences) and the 1000
Plants Project (OneKP or 1KP), as indicated in Table S1.
Phylogenetic analyses. — We initially built four separate matrices for
Papilionoideae, the mimosoid clade, lineages of the former Caesalpinioideae,
and the outgroup taxa. For each, an initial alignment for a subset of taxa was
made using MACSE v1.01b (Ranwez & al., 2011) using default settings, in
order to obtain an alignment that respects the open reading frame (ORF) and
does not allow indels within codons. Running the complete alignment on
MACSE was not possible because it is too computationally intensive. The four
initial alignments were then merged with the MERGE function and additional
sequences added using the --add function in MAFFT v.7 (Katoh & Standley,
2013). The complete matrix was checked by eye and alignments were corrected to ensure that all sequences were aligned with respect to the ORF. An
exception was made for sequences belonging to new sense Caesalpinioideae,
which share a frameshift mutation near the end of the ORF. Two ambiguity
symbols (“ ? ” s) were inserted, disrupting the ORF but ensuring assumed
homology at the nucleotide level.
Aligned matrices were analysed using maximum likelihood and Bayesian
inference. Initial analyses were implemented with RAxML v.8.0 (Stamatakis,
2014) using the GTRGAMMA model with 100 bootstrap replicates to check
for problematic sequences and to ensure that shorter incomplete gene region
sequences did not lead to spurious phylogenetic relationships (e.g., grouping
together of shorter sequences). We used PartitionFinder v.2 (Lanfear & al.,
2012) to determine whether to partition codons separately or not. The program
favoured a single partition for all three codon positions together. Complete
analyses of the final matrix were implemented using a maximum likelihood
approach analysed with RAxML, using the GTRGAMMA model, and support was assessed through 1000 rapid bootstrap replicates. The Bayesian
analyses were implemented in PhyloBayes-MPI v.1.5a (Lartillot & al., 2009),
with the GTR model, running two chains until they reached convergence, as
determined with Tracer v.1.6 (Rambaut & al., 2014). The two chains were
run for a total of 25,891 and 25,512 cycles, and the majority-rule consensus
tree produced by the program bpcomp (included in the PhyloBayes package)
was based on 1075 posterior trees sampled from both chains. A total of 20
accessions were pruned from all trees post-analysis. These were duplicate
accessions or problematically vouchered accessions that were not discovered
until after running the final analyses. We decided to prune these to ensure
that the phylogenetic trees are as clean as possible for potential downstream
comparative analyses. The final RAxML and PhyloBayes analyses were conducted on the Cipres Portal (Miller & al., 2010) and on the ScienceCloud of
the University of Zurich, respectively.
Appendix 2. Glossary of some morphological terms used in Table 1, the key and subfamily descriptions. Illustrations of some key traits are provided in Figs.
12 & 13.
Anthotaxy (inflorescence) – the arrangement of flowers along the inflorescence axis.
Bipinnate (leaves) – a twice pinnately compound leaf, in which leaflets are
arranged in pinnae along the main leaf axis (rachis) (Fig. 12C).
Crescent shaped (hilum) – a U- or V-shaped hilum.
Craspedium (fruit) – an indehiscent fruit that breaks apart, either with the
valves separating as a single unit, or into one-seeded segments (articles),
but leaves the sutures as a persistent margin (the replum) (Fig. 13A).
Drupaceous (fruit) – here used to refer to true drupes and similar fruits. A
drupe is an indehiscent fruit with an outer fleshy part surrounding the
pyrene (“stone”) of hardened endocarp (Fig. 13C).
Exstipellate (leaves) – a leaf with no stipels at the leaflet bases.
Hilum – a scar left on the seed coat from its attachment by a funicle to the
placenta. In subfam. Papilionoideae, the hilum is elongate and split
lengthwise by a hilar groove and the hilar region is usually provided
with a lens (Fig. 13J & K). In the other subfamilies, the hilum is circular,
elliptic, punctiform or crescent shaped and can occur apically, subapically or laterally.
Imparipinnate (leaves) – a pinnately compound leaf (with a rachis) with a
single terminal leaflet (= odd pinnate) (Fig. 12B).
Lens (seed) – a mound situated near the hilum, usually located opposite the
micropyle with the hilum between both structures; an area of weakness
where water initially penetrates the seed prior to germination (Fig. 13J
& K).
Loment (fruit) – a jointed indehiscent fruit (common in legumes) that breaks
apart in one-seeded segments (articles) (Fig. 13B).
Overgrown (seed) – a seed that enlarges and fills the seed-cavity of the pod
without differentiation of the testa and thus the growth is limited by the
size of the pod (Fig. 13F).
Palmate (leaves) – a leaf in which leaflets arise from the apex of the petiole
(i.e., there is no leaf rachis), as fingers originate from the palm of a
hand; in legumes used for such leaves with 4 or more leaflets (i.e., not
for digitately trifoliolate leaves) (Fig. 12D).
Paraphyllidium, plural paraphyllidia (leaves) – reduced leaflets situated
at the base of a pinna-rachis, immediately contiguous to its pulvinus
(Fig. 12F).
Paripinnate (leaves) – a pinnately compound leaf (with a rachis) with a pair
of opposite terminal leaflets (= even pinnate) (Fig. 12A).
Pleurogram (seed) – a fracture line in a seed exotestal palisade leaving a
U- or O-shape on both seed faces (Fig. 13G & H).
Prickles (mechanical defense) – extensions of the plant surface (cortex and
epidermis) with sharp, stiff ends; the prickles detachable without tearing
the organ which they protect.
Pseudopleurogram (seed) – a coloured line on the seed surface but this not
resulting from a break in exotestal palisade (i.e., not a fracture line)
(Fig. 13I).
Pseudoraceme (inflorescence) – a compound raceme in which each bract
subtends two or more flowers in highly condensed lateral axes (Fig.
12G & H).
Samaroid (fruit) – here used to refer to true samaras and similar fruits. A true
samara is a dry, indehiscent, winged fruit, the flattened wing derived
from the ovary wall and usually longer than the seed-bearing part; in
samaroids the wing can encircle the seed chamber (Fig. 13D & E).
Spathaceous (calyx) – a bilaterally symmetrical calyx in which all sepals
are unilaterally joined, usually splitting along one line of weakness at
flower anthesis.
Spines (mechanical defense) – modified leaves, stipules, branches, or parts of
leaves with sharp, stiff ends; always with a vascular origin.
Stipel (leaves) – a stipule-like appendage at the base of a leaflet (Fig. 12E).
Stipellate (leaves) – a leaf with leaflets provided with stipels (Fig. 12E).
Synandrium (androecium) – an androecium in which the stamens are fused
both by the filaments and anthers.
Thyrse (inflorescence) – a panicle composed of cymose lateral units (Fig.
12I & J).
Thyrsoid (inflorescence) – like a thyrse.
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Fig. 12 (Appendix 2). Leaves and inflorescences. A, A paripinniate leaf of Goniorrhachis marginata Taub. (Detarioideae); B, An imparipinnate leaf
of Luetzelburgia bahiensis Yakovlev (Papilionoideae); C, A bipinnate leaf of Pityrocarpa moniliformis (Benth.) Luckow & R.W.Jobson (Caesalpinioideae, mimosoid clade); D, A palmately compound leaf of Zornia myriadena Benth. (Papilionoideae); E, A pinnately trifoliolate leaf of Centrosema
arenarium Benth. (Papilionoideae) highlighting the stipels at the base of leaflets (inset); F, A bipinnate leaf of Mimosa tenuiflora (Willd.) Poir.
(Caesalpinioideae, mimosoid clade) showing a pair of paraphyllidia near the base of the pinna (inset); G, A pseudoraceme of Deguelia nitidula
(Benth.) A.M.G.Azevedo & R.A.Camargo (Papilionoideae) with condensed multiflorous lateral axes; H, Part of a pseudoraceme of Macroptilium
bracteatum (Nees & Mart.) Maréchal & Baudet (Papilionoideae) with biflorous lateral axis; I & J, Thyrsoid inflorescences of Apuleia leiocarpa
(Vogel) J.F.Macbr. (I, Dialioideae) and Zenia insignis Chun (J, Dialioideae). — lfl, leaflet; p, petiole; pn, pinna; prf, paraphyllidium; r, leaf rachis;
stp, stipel. — Photos: G, Luciano P. de Queiroz; H & I, Domingos Cardoso; J, Shijin Li.
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Fig. 13 (Appendix 2). Fruits and seeds. A, A craspedium of Mimosa irrigua Barneby (Caesalpinioideae, mimosoid clade) showing the one-seeded
segments (articles) and the persistent marginal replum (r); B, A loment of Aeschynomene martii Benth. showing the one-seeded segments but no
persistent replum; C, A drupe of Andira humilis Mart. ex Benth.; D & E, Two kinds of samaroid fruits (wings, w): D, Dalbergia nigra (Vell.) Allemão
ex Benth. (Papilionoideae) and E, Luetzelburgia andrade-limae H.C.Lima (Papilionoideae); F, Indehiscent fruit of Dioclea edulis Kuhlm. split
lengthwise to show the overgrown seeds; G & H, Seeds of Caesalpinioideae legumes (mimosoid clade): G, Adenanthera pavonina L. and H, Leucaena
leucocephala (Lam.) De Wit showing the pleurogram (pg); I, Seed of Tamarindus indica L. (Detarioideae) showing the pseudopleurogram (ps);
J–K, Seeds of the common bean: J, Phaseolus vulgaris L. and K, Erythrina velutina Willd. highlighting the major features of Papilionoideae seeds
(insets), with an elongate hilum (h) split lengthwise by a hilar groove (hg) and bearing the micropyle (m) and the lens (l) at the opposite poles of
the hilar region. — Photos: A & E, Domingos Cardoso; B–D & G–K, Luciano P. de Queiroz; F, Alex Popovkin.
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