A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny

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.

Full text

LPWG • Phylogeny and classification of the Leguminosae
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
A new subfamily classification of the Leguminosae based on
a taxonomically comprehensive phylogeny
The Legume Phylogeny Working Group (LPWG)

        

   matK

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* Tani a 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* Mart in F. Wojciechowski,49§* Tetsukazu Yahara,39* Tingshuang Yi54 & Erin Zimmerman2
  Department of Plant Science, University of Tehran, Iran
  Institut de Recherche en Biologie Végétale and Département de Sciences Biologiques, Université de Montréal, Canada
3 Department of Biology, New Mexico State University, Las Cruces, U.S.A.
4 Royal Botanic Gardens, Kew, Richmond, Surrey, U.K.
5 Departamento Ciências Biológicas, Universidade Estadual de Feira de Santana, Brazil
6 Departamento de Biologia Vegetal, Universidade Estadual de Campinas, Brazil
  Department of Biodiversity and Conservation Biology, University of the Western Cape, Cape Town, South Africa
8 Departamento de Botânica, Universidade Federal de São Carlos, Brazil
9 School of BioSciences, University of Melbourne, Australia  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.

Version of Record
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
45
 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
30 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
33 Plant Sciences & Plant Pathology, Montana State University, Bozeman, Montana, U.S.A.
34 South African National Biodiversity Institute, Silverton, South Africa  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
 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.
 Plant Sciences and Biodiversity, Royal Botanic Gardens Victoria, Melbourne, Australia
 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
 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
 Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York, U.S.A.
 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
 Anne Bruneau, anne.bruneau@umontreal.ca
DOI   .3
Abstract

  
matK
 3696
matK






 
              
      
 
                             




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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
46
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. 
A general system of classif ication that is reasonably natural,
mnemonic and traditional is likely to be most useful for most pur-
poses. 
Nutritious seeds for a sustainable future —    


INTRODUCTION
     
          
     

    


     

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   
       
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        
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


 


       
 
 



 







   


 




  
Keywords

matK
Supplementary Material
 matK



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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae








 
  

 



       




 

THE NEW SUBFAMILY CLASSIFICATION
    

 
   
   



    
   

   





  

     

     




  
 







  

  
   

  
Prodromus     




  


Families
of flowering plants  Genera of flowering plants

   



Advances in legume systematics 


   


 

    




    
    






    
         
   






     

        
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TAX ON
66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
48
Fig. .
A,
       matK3696      698 5 
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 matK

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                              
  

 
B,  
matK
 
Fig. .  

matK

  
 




Papilionoideae
(445/503 genera, 2316/ca. 14,000 species)
Caesalpinioideae (incl. mimosoid clade)
(146/148 genera, 937/ca. 4400 species)
Dialioideae
(15/17 genera, 19/ca. 85 species)
Detarioideae
(79/84 genera, 327/ca. 760 species)
Cercidoideae
(12/12 genera, 96/ca. 335 species)
Duparquetioideae
(1/1 genus, 1/1 species)
A.
B.
0.06
mimosoid clade
6.0
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
49Version of Record
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
50
     




Advances in legume
systematics
 


      
         
    
    
  
  
  

      



 
    
       
    

  
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


 
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


   
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 
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  
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matK
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
   


matK
 


 

   
       
   
   
  

     


Duparquetia


 
        

  
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 
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
 
 



   




         
   
  
   








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Fig. . A– F, G, H–L, 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

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Fig. .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. — 

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Fig. . 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. —    

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Fig. .   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 amplis-
simaL, Albizia grandibracteata. —   –  –

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Fig. . 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. — – G–
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Fig. .A, Uleanthus erythrinoidesB, Cadia purpureaC, Sophora 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. —  

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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
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Fig. . A, Chorizema glycinifoliumB, Bossiaea walkeriC, Mucuna giganteaD, Chadsia longidentataE, Canavalia brasilien-
sisF, Erythrina velutinaG, Gliricidia robustaH, Poissonia weberbaueriI, Anthyllis montanaJ, Astragalus uniflorusK, Trifolium rubensL,
Pisum sativumbiflorum. — . d e

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66   : 44–

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Fig. .  A,B,C,D & E,F–L,A, Griffonia
physocarpaB, Duparquetia orchidaceaC, Dialium guianenseD, Brodriguesia santosiiE, Berlinia razziferaF,
Eligmocarpus cynometroidesG, Heteroflorum sclerocarpumH, Erythrostemon coccineusI, Entada polystachyaJ, Prosopis feroxK, Mimosa
townsendiiL, Cojoba arborea. — 

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66   : 44–

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Fig. .A & B,C–L,A, Abarema jupunbaB, Inga feuilleiC, Swartzia parvipetalaD,
Andira micrantha
E,
Crotalaria stipularia
F,
Pterocarpus angolensis
G,
Dalbergia lemurica
H,
Machaerium millei
I,
Carmichaelia . aligera
J, Erythrina madagascariensisK, Piscidia grandifoliaL, Phaseolus— 

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  
  

         
 
  
    
 
    
 
 


           
 
  
  












 
Tab le . 


     



























herbs, or twining
vines with tendrils

 
 Often present 




  Often present on
the petiole and / or
on the primary
and secondary
rachises, usually
between pinnae or
leaflet pairs,



oc-
casionally present
on stipules, stipels,
bracts, or swollen
and nectar- secret-
ing peduncles or
sepals

 Intrapetiolar
(i.e., somewhere
between the petiole
and the axillary
bud) and then free,
valvate and con-
nected by chaffy
hairs, or fused,
either partly (only
at base) or entirely,


 




very rarely
interpetiolar

Unifoliolate or
bifoliolate
Usually paripin-
nate

 Usually imparip-
innate, 



bi-
pinnate

mostly paripinnate




Mostly pari- or
imparipinnate
or palmately
compound,

trifoliolate,


Version of Record
TAX ON
66   : 44–

LPWG • Phylogeny and classification of the Leguminosae


    






  
           

  



 


  


  

 


    
      International Code of
Nomenclature for algae, fungi and plants Melbourne Code

Tab le . 
     
 























  Branched, thyrsoid
inflorescences,














  Large 
frequently petaloid,
valvate, imbricate
or partially fused
or partly fused
with the hypan-
thium, partially or
completely enclos-
ing the bud
   






some-
times papilionate






never papilionate


never
papilionate



sometimes
papilionate









some-
times papilionate




usually
papilionate,



 





 





cupular


Version of Record
TAX ON
66   : 44–

LPWG • Phylogeny and classification of the Leguminosae

Tab le . 
     









4, unequal, free, the
abaxial and adaxial
sepals cucullate
and sepaloid, the
laterals petaloid













 

imbri-
cate, the adaxial
petal innermost



imbricate,
the adaxial petal
generally outer-
most,





imbricate,
the adaxial petal
outermost



all 5
petals with stalked
glands along their
margins

im-
bricate, the adaxial
petal innermost, 



valvate
or imbricate,
then adaxial petal
innermost



imbricate, the
adaxial petal
outermost
















4



















  

Filaments usually
connate into a
sheath or tube




















































rarely
in tetrads




asymmet-
rical, one equa-
torial-encircling
ectoaperture with
two equatorial
endoapertures






porate tetrads,
bitetrads or
polyads,





















rarely polycar-
pellate





Version of Record
TAX ON
66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
63
Tab le . 
     





 
























Commonly inde-
hiscent drupaceous
or samaroid,
























 cres-
cent-shaped hilum,

lack-
ing pleurograms,
pseudopleuro-
grams, wing or aril


occasionally
with pseudopleuro-
grams; 



lacking
pleurograms

lacking
pleurograms
Usually with an
open or closed
pleurogram
on both faces,













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
pleurogram
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    
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    
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
64
Tab le  . 
    matK 
.
CERCIDOIDEAEAdenolobus  Barklya  Bauhinia  Bre-
nierea  Cercis  Gigasiphon  Griffonia  Lysiphyllum   Phanera   Piliostigma  Schnella
 Tylo sem a  
Recent synonym: Lasiobema Phanera  
DETARIOIDEAEAfzelia  Amherstia   Annea  Anthonotha  Aphano-
calyx  Augouardia  Baikiaea  Barnebydendron  Berlinia    Bikinia  *Brachycylix
 Brachystegia  Brandzeia  Brodriguesia  Brownea  Browneopsis  Colophospermum
  Copaifera  Crudia  Cryptosepalum  Cynometra  Daniellia  Detarium  Dicymbe 
 Didelotia  Ecuadendron  Elizabetha  Endertia  Englerodendron   Eperua
 Eurypetalum  Gabonius  Gilbertiodendron  Gilletiodendron     Goniorrhachis 
Gossweilerodendron  Guibourtia   Hardwickia  Heterostemon  Humboldtia  Hylodendron  Hymenaea 
Hymenostegia  Icuria  Intsia  Isoberlinia  Isomacrolobium  Julbernardia
 Kingiodendron  Lebruniodendron  Leonardoxa  *Leucostegane  Librevillea  Loesenera 
Lysidice  Macrolobium  Maniltoa  *Michelsonia  *Micklethwaitia   Microberlinia 
Neoapaloxylon  Neochevalierodendron  Normandiodendron  Oddoniodendron   Oxystigma 
Paloue  Paloveopsis  Paramacrolobium  Peltogyne  Plagiosiphon   Polystemonanthus  Prioria
 *Pseudomacrolobium  Saraca  Schotia  Scorodophloeus  Sindora  Sindoropsis  Stemonocoleus
 Talbotiella  Tamarindus  Tessmannia  Tetraberlinia  Zenkerella 
Recent synonym: Pellegriniodendron Gilbertiodendron  
DUPARQUETIOIDEAEDuparquetia 
DIALIOIDEAE*Androcalymma  Apuleia  Baudouinia  Dialium  Dicorynia   Diste-
monanthus  Eligmocarpus  Kalappia   Koompassia  Labichea  Martiodendron
 Mendoravia  Petalostylis  Poeppigia  Storckiella  *Uittienia  Zenia 
CAESALPINIOIDEAEAbarema   Acacia
 Acaciella   Acrocarpus   Adenanthera L. Adenopodia  Afrocalliandra  
Alantsilodendron  Albizia  Amblygonocarpus  Anadenanthera  Arapatiella  Archidendron
 Archidendropsis  Arcoa  Arquita    Aubrevillea Balizia 
       Balsamocarpon  Batesia   Biancaea    Blanchetiodendron   Burkea
 Bussea  Caesalpinia  Calliandra  Calliandropsis   Calpocalyx  Campsiandra 
Cassia  Cathormion  Cedrelinga  Cenostigma   Ceratonia  Chamaecrista  Chidlowia  Chloroleucon
 Cojoba  Colvillea  Conzattia  Cordeauxia  Coulteria  Cylicodiscus
 Delonix  Denisophytum       Desmanthus  Dichrostachys  Dimorphandra  Dinizia 
Diptychandra   Ebenopsis  Elephantorrhiza  Entada  Enterolobium  Erythrophleum 
Erythrostemon  Faidherbia  Falcataria  Fillaeopsis  Gagnebina 
Gelrebia  ; Gleditsia  Guilandina  Gymnocladus  Haematoxylum  Havardia Hererolandia    
; Hesperalbizia   Heteroflorum  Hoffmannseggia  *Hultholia   ; Hydrochorea
 *Indopiptadenia  Inga  Jacqueshuberia  Kanaloa  Lemurodendron 
Lemuropisum  Leucaena  Leucochloron  Libidibia        Lophocarpinia  Lysiloma
 Macrosamanea            Mariosousa   Melanoxylon  Mezoneuron  Microlobius
 Mimosa L. Mimozyganthus  Moldenhawera  Mora  Moullava  Neptunia  Newtonia
 Pachyelasma  Painteria   Parapiptadenia  Pararchidendron  Paraserianthes  Parkia
 Parkinsonia Paubrasilia; Peltophorum  Pentaclethra  Piptadenia 
Piptadeniastrum  Piptadeniopsis  Pithecellobium  Pityrocarpa  Plathymenia  Pomaria
  Prosopidastrum  Prosopis L. Pseudopiptadenia  Pseudoprosopis  Pseudosamanea Pterogyne  
Pterolobium  Recordoxylon  Samanea  Sanjappa    ; Schizolobium   
Schleinitzia  Senegalia  Senna  Serianthes  Sphinga   Stachyothyrsus
 Stenodrepanum  Stryphnodendron  Stuhlmannia    Sympetalandra  Tachigali  Tara  Tet r apleura
 Tetrapterocarpon   Thailentadopsis  Umtiza  Vachellia    Viguieranthus  Vouacapoua 
Wallaceodendron  Xerocladia    Xylia  Zapoteca  Zuccagnia    Zygia      .
Recent synonyms: GuinetiaCalliandra Marmaroxylon Zygia        Poincianella 
ErythrostemonCenostigma  Stahlia Libidibia
PAPILIONOIDEAE  Abrus   Acmispon  Acosmium  Adenocarpus  Adenodolichos
 Adesmia  Aenictophyton   Aeschynomene  Afgekia  Aganope  Airyantha  Akschindlium 
Aldina  Alexa  Alhagi  Alistilus   Almaleea     Alysicarpus   Amburana   
Amicia  Ammodendron  Ammopiptanthus  Ammothamnus  Amorpha  Amphicarpaea    
Amphimas        Amphiodon  Amphithalea  Anagyris  Anarthrophyllum  Ancistrotropis 
Andira  Angylocalyx  Antheroporum  Anthyllis  *Antopetitia  Aotus  Aphyllodium  Apios
 Apoplanesia  Apurimacia  Arachis  Argyrocytisus  Argyrolobium  Arthroclianthus
 Aspalathus  Astragalus  Ateleia  Austrodolichos    Austrosteenisia  Baphia 
    Baphiastrum  Baphiopsis  Baptisia  *Barbieria  Behaimia  Bionia  *Biser-
rula  Bituminaria  Bobgunnia  Bocoa  Bolusafra  Bolusanthus  Bolusia 
Bossiaea  Bowdichia  Bowringia  Brongniartia  Brya        Bryaspis           *Burkilliodendron
 Butea  Cadia  Cajanus  Calicotome  Callerya  Callistachys   Calobota  
Calophaca  Calopogonium  Calpurnia  Camoensia  Camptosema  Campylotropis
 Canavalia  Candolleodendron  Caragana  Carmichaelia  Carrissoa    Cascaronia  
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
65
Castanospermum     Centrolobium  Centrosema  Chadsia  Chaetocalyx  Chapmannia
    Chesneya  Chorizema  Christia  *Chrysoscias  Cicer  Cladrastis  Clathrotropis
 Cleobulia  Clianthus  Clitoria  Clitoriopsis        *Cochlianthus  Cochliasanthus
 Codariocalyx  Collaea  Cologania  Colutea  Condylostylis  Cordyla  Corethrodendron 
Coronilla  Coursetia  Craibia   Cranocarpus  Craspedolobium   Cratylia   *Cristonia
 Crotalaria  Cruddasia  Cullen   Cyamopsis  Cyathostegia  Cyclocarpa  Cyclolo-
bium  Cyclopia   Cymbosema  Cytisophyllum  *Cytisopsis   Cytisus  Dahlstedtia 
Dalbergia  Dalbergiella  Dalea  Dalhousiea  Daviesia  Decorsea      Deguelia  Dendro-
lobium   Dermatophyllum  Derris  *Desmodiastrum  Desmodium  
Dewevrea  Dichilus  Dicraeopetalum  Dillwynia  Dioclea  Diphyllarium  Diphysa  Diplotropis
 Dipogon  Dipteryx  Discolobium  Disynstemon  Dolichopsis  Dolichos  Dorycnium 
*Dorycnopsis  Droogmansia  Dumasia  Dunbaria   Dussia   Dysolobium  
 Ebenus  *Echinospartum  Eleiotis  *Eminia  Endosamara  Eremosparton 
Erichsenia  Erinacea  Eriosema  Erophaca  Errazurizia  Erythrina  Euchilopsis    Euchlora
 *Euchresta   Eutaxia  Eversmannia  Exostyles  Eysenhardtia  *Ezoloba 
   Fairchildia      Fiebrigiella  Fissicalyx  Flemingia    Ford ia  Galactia
      Galega  Gastrolobium  Geissaspis  Genista  Genistidium  Geoffroea 
Gliricidia  Glycine  Glycyrrhiza  Gompholobium  *Gonocytisus  Goodia Grazielodendron 
*Greuteria      Gueldenstaedtia  Guianodendron  Halimodendron 
 Hammatolobium   Hanslia   Haplormosia  Hardenbergia  Harleyodendron  Harpalyce 
     Haymondia       Hebestigma  Hedysarum  Hegnera  Helicotropis  *Herpyza 
Hesperolaburnum  Hesperothamnus  Hippocrepis   Hoita  Holocalyx  *Hosackia  Hovea
 Humularia  Hylodesmum   Hymenocarpos  Hymenolobium  Hypocalyptus
 Indigastrum  Indigofera  Inocarpus  Isotropis  Jacksonia 
*Kebirita       Kennedia   Kotschya  Kummerowia  Kunstleria  Lablab   Laburnum  
Lackeya   Ladeania  Lamprolobium  Lathyrus  Latrobea  *Lebeckia
 Lecointea  *Lembotropis  Lennea  Lens  Leobordea  Leptoderris  *Leptodesmia   
 Leptolobium    Leptosema  Leptospron  Lespedeza  Lessertia  Leucomphalos
 Limadendron    Liparia  *Listia    Lonchocarpus  Lotononis 
Lotus  Luetzelburgia  Lupinus  Luzonia  Maackia      Machaerium  Macropsychanthus  Macrop -
tilium  Macrotyloma   Maraniona    Marina  Mastersia
 Mecopus  Medicago  *Meizotropis    Melilotus  Melliniella  Melolobium  Microcharis 
Mildbraediodendron  Millettia  Mirbelia  Monarthrocarpus   Monopteryx   *Montigena
 Mucuna  Muellera  Muelleranthus  Mundulea  Myrocarpus  Myrospermum  Myroxylon
 Mysanthus  *Neocollettia  *Neoharmsia      Neonotonia  Neorautanenia  Neoru-
dolphia  Nephrodesmus  Nesphostylis    Neustanthus  Nissolia  Nogra  Oberholzeria 
    Ohwia  Olneya   Onobrychis  Ononis  Ophrestia  Orbexilum
 *Oreophysa  Ormocarpopsis      Ormocarpum  Ormosia  Ornithopus  Orphanodendron
   Oryxis  *Ostryocarpus   Otholobium  Otoptera  *Ototropis 
*Ottleya   *Ougeinia  Oxylobium  Oxyrhynchus  Oxytropis  Pachyrhizus  Panurea
 Paracalyx  *Paragoodia  Paramachaerium  Paratephrosia  Parochetus 
Parryella   Pearsonia  Pediomelum  Periandra  Pericopsis  Petaladenium
 Peteria  Petteria  Phaseolus  Philenoptera  *Phylacium  Phyllodium   Phyllolobium
 Phyllota  Phylloxylon  Physostigma  Pickeringia   Pictetia  Piptanthus 
Piscidia  Pisum   Plagiocarpus  Platycelyphium   Plat ycyamus  Platylobium  Platymiscium    Platypodium
  Platysepalum  Podalyria  Podlechiella   *Podocytisus    Podolobium
 *Podolotus  Poecilanthe  Poiretia    *Poissonia  Poitea   Polhillia  Pongamiopsis
 Pseudarthria    Pseudeminia   Pseudoeriosema  *Pseudolotus  Pseudovigna        
Psophocarpus  Psoralea  *Psoralidium  Psorothamnus  Pterocarpus  Pterodon    Ptycholobium 
*Ptychosema  Pueraria  Pultenaea  Pycnospora      *Pyranthus       Rafnia  Ramirez-
ella  Ramorinoa  Requienia  Retama  Rhodopis  Rhynchosia    *Rhynchotropis  Riedeliella 
Robinia  *Robynsiophyton       *Rothia  Rupertia         *Sakoanala  *Salweenia   *Sarcodum  
Sartoria  Schefflerodendron  Scorpiurus  Securigera  Sellocharis   Sesbania  Shuteria 
 Sigmoidotropis  Sinodolichos    Smirnowia  Smithia  Soemmeringia  Solori  Sophora
 Spartium  Spathionema    Spatholobus  Sphaerolobium  Sphaerophysa  Sphenostylis  Sphinctospermum
 Spirotropis   Spongiocarpella     Staminodianthus  Stauracanthus
 Steinbachiella  Stirtonanthus     *Stonesiella      *Streblorrhiza  Strongylodon
  Strophostyles  Stylosanthes  Styphnolobium  Sulla  Sutherlandia  Swainsona 
Swartzia  Sweetia  *Sylvichadsia    *Syrmatium   Tab aro a  Tadeh agi
 Taralea  Taverniera  Templetonia   Tephrosia  *Teramnus        Tetragonolobus 
Teyleria  Thermopsis    Thinicola  Tibetia           Tipuana  Toxicopueraria 
 Trifidacanthus  Trifolium  Trigonella  Tripodion  Trischidium  Uleanthus  Ulex  Uraria  
Uribea  Urodon    *Vandasina  Vatairea  Vataireopsis  Vatovaea    Vavilovia  
*Verdesmum  Vicia  Vigna  Viminaria  Virgilia  *Vuralia  Waj ir a  Weber-
bauerella  *Wiborgia  *Wiborgiella     Wisteria  Xanthocercis  *Xeroderris  Xiphotheca
 Zollernia  Zornia      Zygocarpum 
Recent synonyms: Barnebyella AstragalusBergeronia  MuelleraCalia  Dermatophyllum
Etaballia PterocarpusMargaritolobiumMuelleraOphiocarpus  AstragalusParaderris   
DerrisPeltiera OrmocarpopsisSpartidium Calobota
Tab le .
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66   : 44–

LPWG • Phylogeny and classification of the Leguminosae
66
     


   












 
 



  

  


INTEGRATING TRIBAL AND CLADE-BASED
CLASSIFICATIONS
    



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