Editing Fabaceae (section)

== Evolution, phylogeny and taxonomy ==

=== Evolution ===

The order Fabales contains around 7.3% of eudicot species and the greatest part of this diversity is contained in just one of the four families that the order contains: Fabaceae. This clade also includes the families [[Polygalaceae]], [[Surianaceae]] and [[Quillajaceae]] and its origins date back 94 to 89 million years, although it started its diversification 79 to 74 million years ago.<ref name="Stevens 2001" /> The Fabaceae diversified during the [[Paleogene]] to become a ubiquitous part of the modern earth's [[Biota (ecology)|biota]], along with many other families belonging to the flowering plants.<ref name="Lewis" /><ref name="Herendeen92">Herendeen, P. S., W. L. Crepet, and D. L. Dilcher. 1992. The fossil history of the Leguminosae: phylogenetic and biogeographic implications. Pages 303 – 316 in Advances in Legume Systematics, part 4, the fossil record (P. S. Herendeen and D .L. Dilcher, eds). Royal Botanic Gardens, Kew, UK.</ref>

The Fabaceae have an abundant and diverse [[fossil]] record, especially for the [[Tertiary]] period. Fossils of flowers, fruit, leaves, wood and [[pollen]] from this period have been found in numerous locations.<ref>{{cite journal |last1=Crepet |first1=W. L. |last2=Taylor |first2=D. W. |title=The Diversification of the Leguminosae: First Fossil Evidence of the Mimosoideae and Papilionoideae |journal=Science |volume=228 |issue=4703 |year=1985 |pages=1087–1089 |issn=0036-8075 |doi=10.1126/science.228.4703.1087 |pmid=17737903 |bibcode=1985Sci...228.1087C |s2cid=19601874}}</ref><ref name="Crepet86">{{cite journal |author1=Crepet, W. L. |author2=D. W. Taylor |year=1986 |title= Primitive mimosoid flowers from the Palaeocene-Eocene and their systematic and evolutionary implications. |journal= American Journal of Botany |volume=73 |issue=4 |pages=548–563 |jstor=2444261 |doi=10.2307/2444261}}</ref><ref name="Crepet92">Crepet, W. L., and P. S. Herendeen. 1992. Papilionoid flowers from the early Eocene of south eastern North America. Pages 43–55 in Advances in Legume Systematics, part 4, the fossil record (P. S. Herendeen and D. L. Dilcher, eds.). Royal Botanic Gardens, Kew, UK.</ref><ref name=autogenerated2>Herendeen, P. S. 1992. The fossil history of Leguminosae from the Eocene of south eastern North America. Pages 85–160 in Advances in Legume Systematics, part 4, the fossil record (Herendeen, P. S., and D. L. Dilcher, eds.). Royal Botanic Gardens, Kew, UK.</ref><ref name="Herendeen01">Herendeen, P. S. 2001. The fossil record of the Leguminosae: recent advances. In Legumes Down Under: the Fourth International Legume conference, Abstracts, 34–35. Australian National University, Canberra, Australia.</ref><ref name=autogenerated3>Herendeen, P. S., and S. Wing. 2001. Papilionoid legume fruits and leaves from the Palaeocene of north western Wyoming. Botany 2001 Abstracts, published by Botanical Society of America (http://www.botany2001.org/).</ref><ref name="Wing">Wing, S. L., F. Herrera, and C. Jaramillo. 2004. A Palaeocene flora from the Cerrajón Formation, Guajíra Peninsula, north eastern Colombia. Pages 146–147 in VII International Organization of Paleobotany Conference Abstracts (21–26 March). Museo Egidio Feruglio, Trelew, Argentina.</ref> The earliest fossils that can be definitively assigned to the Fabaceae appeared in the early [[Palaeocene]] (approximately 65 million years ago).<ref>{{Cite journal |last1=Herendeen |first1=Patrick S. |last2=Cardoso |first2=Domingos B. O. S. |last3=Herrera |first3=Fabiany |last4=Wing |first4=Scott L. |date=January 2022 |title=Fossil papilionoids of the Bowdichia clade (Leguminosae) from the Paleogene of North America |journal=American Journal of Botany |language=en |volume=109 |issue=1 |pages=130–150 |doi=10.1002/ajb2.1808 |issn=0002-9122 |pmc=9306462 |pmid=35014023}}</ref> Representatives of the 3 sub-families traditionally recognised as being members of the Fabaceae – Cesalpinioideae, Papilionoideae and Mimosoideae{{Snd}}as well as members of the large clades within these sub-families{{Snd}}such as the genistoides{{Snd}}have been found in periods later, starting between 55 and 50 million years ago.<ref name="Herendeen92" /> In fact, a wide variety of taxa representing the main lineages in the Fabaceae have been found in the fossil record dating from the middle to the late [[Eocene]], suggesting that the majority of the modern Fabaceae groups were already present and that a broad diversification occurred during this period.<ref name="Herendeen92" /> Therefore, the Fabaceae started their diversification approximately 60 million years ago and the most important clades separated 50 million years ago.<ref name="Bruneau2008b">Bruneau, A., Lewis, G. P., Herendeen, P. S., Schrire, B., & Mercure, M. 2008b. Biogeographic patterns in early-diverging clades of the Leguminosae. Pp. 98–99, in Botany 2008. Botany without Borders. [Botanical Society of America, Abstracts.]</ref> The age of the main Cesalpinioideae clades have been estimated as between 56 and 34 million years and the basal group of the Mimosoideae as 44 ± 2.6 million years.<ref name="Bruneau2008a">{{cite journal |author1=Bruneau, A. |author2=Mercure, M. |author3=Lewis, G. P. |author4= Herendeen, P. S. |name-list-style=amp |year=2008 |title=Phylogenetic patterns and diversification in the caesalpinioid legumes |journal=Canadian Journal of Botany |volume=86 |issue=7 |pages=697–718 |doi=10.1139/B08-058}}</ref><ref name=autogenerated1>{{cite journal |author=Lavin, M., Herendeen, P. S., y Wojciechowski, M. F. |year=2005 |title=Evolutionary Rates Analysis of Leguminosae Implicates a Rapid Diversification of Lineages during the Tertiary |journal=Systematic Biology |volume=54 |issue=4 |pages=575–594 |doi=10.1080/10635150590947131 |pmid=16085576 |last2=Herendeen |last3=Wojciechowski |doi-access=free }}</ref> The division between Mimosoideae and Faboideae is dated as occurring between 59 and 34 million years ago and the basal group of the Faboideae as 58.6 ± 0.2 million years ago.<ref name="Wikström">{{cite journal |last1=Wikstrom |first1=N. |last2=Savolainen |first2=V. |last3=Chase |first3=M. W. |title=Evolution of the angiosperms: calibrating the family tree |journal=Proceedings of the Royal Society B: Biological Sciences |date=2001 |volume=268 |issue=1482 |pages=2211–2220 |doi=10.1098/rspb.2001.1782 |pmid=11674868 |pmc=1088868}}</ref> It has been possible to date the divergence of some of the groups within the Faboideae, even though diversification within each genus was relatively recent. For instance, ''[[Astragalus (plant)|Astragalus]]'' separated from the ''[[Oxytropis]]'' 16 to 12 million years ago. In addition, the separation of the [[aneuploidy|aneuploid]] species of ''Neoastragalus'' started 4 million years ago. ''[[Inga]]'', another genus of the Papilionoideae with approximately 350 species, seems to have diverged in the last 2 million years.<ref name="Wojciechowski03">Wojciechowski, M. F. 2003. Reconstructing the phylogeny of legumes (Leguminosae): An early 21st century perspective. Pp. 5–35, in Klitgaard, B. B. & Bruneau, A. (eds), Advances in Legume Systematics, Part 10, Higher Level Systematics. Royal Botanic Gardens, Kew.</ref><ref name="Wojciechowski04">{{cite journal |last1=Wojciechowski |first1=M. F. |title=Astragalus (Fabaceae): A molecular phylogenetic perspective |journal=Brittonia |date=2005 |volume=57 |issue=4 |pages=382–396 |doi=10.1663/0007-196X(2005)057[0382:AFAMPP]2.0.CO;2 |jstor=4098954 |s2cid=21645067 }}</ref><ref name="Wojciechowski93">{{cite journal |last1=Wojciechowski |first1=M. F. |last2=Sanderson |first2=M. J. |last3=Baldwin |first3=B. G. |last4=Donoghue |first4=M. J. |date=1993 |title=Monophyly of aneuploid ''Astragalus'': Evidence from nuclear ribosomal DNA internal transcribed spacer sequences |journal=American Journal of Botany |volume=80 |issue=6 |pages=711–722 |jstor=2445441 |doi=10.2307/2445441}}</ref><ref name="Wojciechowski06">Wojciechowski, Martin F., Johanna Mahn, and Bruce Jones. 2006. Fabaceae. legumes. Version 14 June 2006. [http://tolweb.org/Fabaceae/21093/2006.06.14 The Tree of Life Web Project], http://tolweb.org/</ref>

It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along the [[Tethys Ocean|Tethys seaway]] during the [[Palaeogene]] Period.<ref name="Schrire2">{{cite book |last1=Schrire |first1=B. D. |last2=Lewis |first2=G. P. |last3=Lavin |first3=M. |editor1-first=G |editor1-last=Lewis |editor2-first=G. |editor2-last=Schrire |editor3-first=B. |editor3-last=Mackinder |editor4-first=M. |editor4-last=Lock |title=Legumes of the world |chapter-url=http://www.kewbooks.com/asps/ShowDetails.asp?id=506 |year=2005 |publisher=Royal Botanic Gardens |location=Kew, England |isbn=978-1-900347-80-8 |pages=21–54 |chapter=Biogeography of the Leguminosae |access-date=8 July 2010 |archive-date=2 February 2014 |archive-url=https://web.archive.org/web/20140202160032/http://www.kewbooks.com/asps/ShowDetails.asp?id=506 |url-status=dead }}</ref><ref name="Schrire1">{{cite book |last1=Schrire |first1=B. D. |last2=Lavin |first2=M. |last3=Lewis |first3=G. P. |editor1-first=I |editor1-last=Friis |editor2-first=H. |editor2-last=Balslev |title=Plant diversity and complexity patterns: local, regional and global dimensions |series=Biologiske Skrifter |volume=55 |year=2005 |publisher=Special-Trykkeriet Viborg A/S |location=Viborg, Denmark |isbn=978-87-7304-304-2 |pages=375–422 |chapter=Global distribution patterns of the Leguminosae: insights from recent phylogenies}}</ref> However, others contend that [[Africa]] (or even the [[Americas]]) cannot yet be ruled out as the origin of the family.<ref>{{cite journal |doi=10.1111/j.1095-8339.2010.01044.x |title=Detarieae ''sensu lato'' (Fabaceae) from the Late Oligocene (27.23 Ma) Guang River flora of north-western Ethiopia |year=2010 |last1=Pan |first1=Aaron D. |last2=Jacobs |first2=Bonnie F. |last3=Herendeen |first3=Patrick S. |journal=Botanical Journal of the Linnean Society |volume=163 |pages=44–54 |doi-access=free }}</ref><ref>{{cite journal |doi=10.1104/pp.102.018150 |title=The Rest of the Iceberg. Legume Diversity and Evolution in a Phylogenetic Context |year=2003 |last1=Doyle |first1=J. J. |journal=Plant Physiology |volume=131 |issue=3 |pages=900–10 |pmid=12644643 |last2=Luckow |first2=M. A. |pmc=1540290}}</ref>

The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event.<ref>{{cite journal |doi=10.1007/s00018-011-0651-4 |title=Function and evolution of nodulation genes in legumes |year=2011 |last1=Yokota |first1=Keisuke |last2=Hayashi |first2=Makoto |s2cid=13154916 |journal=Cellular and Molecular Life Sciences |volume=68 |issue=8 |pages=1341–51 |pmid=21380559 |pmc=11114672 }}</ref> Several different pathways have been implicated as donating duplicated genes to the pathways need for nodulation. The main donors to the pathway were the genes associated with the arbuscular mycorrhiza symbiosis genes, the pollen tube formation genes and the haemoglobin genes. One of the main genes shown to be shared between the arbuscular mycorrhiza pathway and the nodulation pathway is SYMRK and it is involved in the plant-bacterial recognition.<ref>{{cite journal |doi=10.1371/journal.pbio.0060068 |title=Functional Adaptation of a Plant Receptor- Kinase Paved the Way for the Evolution of Intracellular Root Symbioses with Bacteria |year=2008 |last1=Markmann |first1=Katharina |last2=Giczey |first2=Gábor |last3=Parniske |first3=Martin |journal=PLOS Biology |volume=6 |issue=3 |pages=e68 |pmid=18318603 |pmc=2270324 |doi-access=free }}</ref> The pollen tube growth is similar to the infection thread development in that infection threads grow in a polar manner that is similar to a pollen tubes polar growth towards the ovules. Both pathways include the same type of enzymes, pectin-degrading cell wall enzymes.<ref>{{cite journal |doi=10.1111/j.1365-313X.2004.02155.x |title=From pollen tubes to infection threads: Recruitment of Medicago floral pectic genes for symbiosis |year=2004 |last1=Rodríguez-Llorente |first1=Ignacio D. |last2=Pérez-Hormaeche |first2=Javier |last3=Mounadi |first3=Kaoutar El |last4=Dary |first4=Mohammed |last5=Caviedes |first5=Miguel A. |last6=Cosson |first6=Viviane |last7=Kondorosi |first7=Adam |last8=Ratet |first8=Pascal |last9=Palomares |first9=Antonio J. |journal=The Plant Journal |volume=39 |issue=4 |pages=587–98 |pmid=15272876 |doi-access=free }}</ref> The enzymes needed to reduce nitrogen, nitrogenases, require a substantial input of ATP but at the same time are sensitive to free oxygen. To meet the requirements of this paradoxical situation, the plants express a type of haemoglobin called leghaemoglobin that is believed to be recruited after a duplication event.<ref>{{cite journal |doi=10.1016/j.cub.2005.03.007 |title=Legume Haemoglobins: Symbiotic Nitrogen Fixation Needs Bloody Nodules |year=2005 |last1=Downie |first1=J. Allan |journal=Current Biology |volume=15 |issue=6 |pages=R196–8 |pmid=15797009 |s2cid=17152647 |doi-access=free |bibcode=2005CBio...15.R196D }}</ref> These three genetic pathways are believed to be part of a gene duplication event then recruited to work in nodulation.

=== Phylogeny and taxonomy ===

==== Phylogeny ====

The [[Phylogenetics|phylogeny]] of the legumes has been the object of many studies by research groups from around the world. These studies have used morphology, [[DNA]] data (the [[chloroplast]] [[intron]] ''trnL'', the chloroplast [[gene]]s ''rbcL'' and ''matK'', or the ribosomal spacers ''ITS'') and [[Cladistics|cladistic analysis]] in order to investigate the relationships between the family's different lineages. Fabaceae is consistently recovered as [[Monophyly|monophyletic]].<ref name="tolweb">{{cite web |url=http://tolweb.org/Fabaceae/21093/2006.06.14 |title=Fabaceae |author1=Martin F. Wojciechowski |author2=Johanna Mahn |author3=Bruce Jones |year=2006 |work=The Tree of Life Web Project }}</ref> The studies further confirmed that the traditional subfamilies Mimosoideae and Papilionoideae were each [[Monophyly|monophyletic]] but both were nested within the paraphyletic subfamily Caesalpinioideae.<ref name="wojciechowski2004">{{cite journal |doi=10.3732/ajb.91.11.1846 |title=A phylogeny of legumes (Leguminosae) based on analysis of the plastid matK gene resolves many well-supported sub clades within the family |year=2014 |author1=Wojciechowski, M. F. |author2=Lavin, M. |author3=Sanderson, M. J. |journal=American Journal of Botany |volume=91 |pages=1846–62 |issue=11 |pmid=21652332 |doi-access=free }}</ref><ref name="tolweb" /> All the different approaches yielded similar results regarding the relationships between the family's main clades.<ref name="Stevens 2001" /><ref name="Käss">{{cite journal |last1=Käss |first1=E. |last2=Wink |first2=M. |year=1996 |title=Molecular evolution of the Leguminosae: phylogeny of the three subfamilies based on ''rbcL'' sequences |journal=Biochemical Systematics and Ecology |volume=24 |issue=5 |pages=365–378 |doi=10.1016/0305-1978(96)00032-4 |bibcode=1996BioSE..24..365K }}</ref><ref name="Käss2">{{cite journal |last1=Käss |first1=E. |last2=Wink |first2=M. |year=1997 |title=Phylogenetic relationships in the Papilionoideae (Family Leguminosae) based on nucleotide sequences of cpDNA (''rbcL'') and ncDNA (ITS1 and 2) |journal=[[Molecular Phylogenetics and Evolution|Mol. Phylogenet. Evol.]] |volume=8 |issue=1 |pages=65–88 |pmid=9242596 |doi=10.1006/mpev.1997.0410 |bibcode=1997MolPE...8...65K }}</ref><ref name="Doyle">{{cite journal |last=Doyle |first=Jeff J. |last2=Doyle |first2=Jane L. |last3=Ballenger |first3=Julie A. |last4=Dickson |first4=Elizabeth E. |last5=Kajita |first5=Tadashi |last6=Ohashi |first6=Hiroyoshi |year=1997 |title=A phylogeny of the chloroplast gene ''rbcL'' in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation |journal=[[American Journal of Botany]] |volume=84 |issue=4 |pages=541–554 |pmid=21708606 |doi=10.2307/2446030 |jstor=2446030 |doi-access=free }}</ref><ref name="Lavin">{{cite journal |last1=Lavin |first1=M. |last2=Doyle |first2=J. J. |last3=Palmer |first3=J. D. |year=1990 |title=Evolutionary significance of the loss of the chloroplast-DNA inverted repeat in the Leguminosae subfamily Papilionoideae |journal=[[Evolution (journal)|Evolution]] |volume=44 |issue=2 |pages=390–402 |jstor=2409416 |doi=10.2307/2409416 |pmid=28564377 |hdl=2027.42/137404 |url=https://deepblue.lib.umich.edu/bitstream/2027.42/137404/1/evo05207.pdf |hdl-access=free }}</ref><ref name="Sanderson">{{cite journal |last1=Sanderson |first1=M. J. |last2=Wojciechowski |first2=M. F. |year=1996 |title=Diversification rates in a temperate legume clade: are there "so many species" of ''Astragalus'' (Fabaceae)? |journal=[[American Journal of Botany|Am. J. Bot.]] |volume=83 |issue=11 |pages=1488–1502 |jstor=2446103 |doi=10.2307/2446103}}</ref><ref name="Chappill">{{cite book |author=Chappill, J. A. |year=1995 |chapter=Cladistic analysis of the Leguminosae: the development of an explicit hypothesis |pages=1–10 |title=Advances in Legume Systematics, Part 7: Phylogeny |editor1=Crisp, M. D. |editor2=Doyle, J. J. |publisher=Royal Botanic Gardens, Kew, UK |isbn=9780947643799}}</ref><ref>{{cite journal |last=Bruneau |first=Anne |last2=Mercure |first2=Marjorie |last3=Lewis |first3=Gwilym P. |last4=Herendeen |first4=Patrick S. |year=2008 |title=Phylogenetic patterns and diversification in the caesalpinioid legumes |journal=[[Botany (journal)|Botany]] |volume=86 |issue=7 |pages=697–718 |doi=10.1139/B08-058}}</ref><ref>{{cite journal |last=Cardoso |first=D. |last2=Pennington |first2=R. T. |last3=de Queiroz |first3=L. P. |last4=Boatwright |first4=J. S. |last5=Van Wyk |first5=B.-E. |last6=Wojciechowski |first6=M. F. |last7=Lavin |first7=M. |year=2013 |title=Reconstructing the deep-branching relationships of the papilionoid legumes |journal=S. Afr. J. Bot. |volume=89 |pages=58–75 |doi=10.1016/j.sajb.2013.05.001 |doi-access=free |hdl=10566/3193 |hdl-access=free }}</ref>{{excessive citations inline |date=December 2024}} Following extensive discussion in the legume phylogenetics community, the Legume Phylogeny Working Group reclassified Fabaceae into six subfamilies, which necessitated the segregation of four new subfamilies from Caesalpinioideae and merging Caesapinioideae ''sensu stricto'' with the former subfamily Mimosoideae.<ref name="6subfamilies">{{cite journal |author=The Legume Phylogeny Working Group (LPWG). |year=2017 |title=A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny |journal=[[Taxon (journal)|Taxon]] |volume=66 |issue=1 |pages=44–77 |doi=10.12705/661.3 |doi-access=free |hdl=10568/90658 |hdl-access=free }}</ref><ref>{{cite journal |last=Koenen |first=Erik J. M. |last2=Ojeda |first2=Dario I. |last3=Steeves |first3=Royce |last4=Migliore |first4=Jérémy |last5=Bakker |first5=Freek T. |last6=Wieringa |first6=Jan J. |last7=Kidner |first7=Catherine |last8=Hardy |first8=Olivier J. |last9=Pennington |first9=R. Toby |last10=Bruneau |first10=Anne |last11=Hughes |first11=Colin E. |year=2019 |title=Large-scale genomic sequence data resolve the deepest divergences in the legume phylogeny and support a near-simultaneous evolutionary origin of all six subfamilies |journal=New Phytologist |volume=225 |issue=3 |pages=1355–1369 |doi=10.1111/nph.16290 |pmid=31665814 |pmc=6972672 |doi-access=free }}</ref> The exact branching order of the different subfamilies is still unresolved.<ref>{{cite journal |last=Zhang |first=Rong |last2=Wang |first2=Yin-Huan |last3=Jin |first3=Jian-Jun |last4=Stull |first4=Gregory W |last5=Bruneau |first5=Anne |last6=Cardoso |first6=Domingos |last7=De Queiroz |first7=Luciano Paganucci |last8=Moore |first8=Michael J |last9=Zhang |first9=Shu-Dong |last10=Chen |first10=Si-Yun |last11=Wang |first11=Jian |last12=Li |first12=De-Zhu |last13=Yi |first13=Ting-Shuang |year=2020 |title=Exploration of Plastid Phylogenomic Conflict Yields New Insights into the Deep Relationships of Leguminosae |journal=Syst. Biol. |volume=69 |issue=4 |pages=613–622 |doi=10.1093/sysbio/syaa013 |pmid=32065640 |pmc=7302050 }}</ref>

{{Clade |style=line-height:75%;
 |label1=[[Fabales]]
 |1={{clade
  |1=[[Polygalaceae]] ([[Outgroup (cladistics)|outgroup]])
  |2=[[Surianaceae]] (outgroup)
  |3={{clade
   |1=[[Quillajaceae]] (outgroup)
   |label2='''Fabaceae'''
   |2={{clade
    |1={{clade
     |1=[[Cercidoideae]]
     |2=[[Detarioideae]]
     }}
    |2={{clade
     |1=[[Duparquetioideae]]
     |2={{clade
      |1=[[Dialioideae]]
      |2={{clade
       |1=[[Caesalpinioideae]]
       |2=[[Faboideae]]
       }}
      }}
     }}
    }}
   }}
  }}
}}

==== Taxonomy ====

The Fabaceae are placed in the order [[Fabales]] according to most taxonomic systems, including the [[APG III system]].<ref name=APGIII2009 /> The family now includes six subfamilies:<ref name="6subfamilies"/>
* [[Cercidoideae]]: 12 genera and ~335 species. Mainly tropical. ''[[Bauhinia]]'', ''[[Cercis]]''.
* [[Detarioideae]]: 84 genera and ~760 species. Mainly tropical. ''[[Amherstia]]'', ''[[Detarium]]'', ''[[Tamarindus]]''.
* [[Duparquetioideae]]: 1 genus and 1 species. West and Central Africa. ''[[Duparquetia]]''.
* [[Dialioideae]]: 17 genera and ~85 species. Widespread throughout the tropics. ''[[Dialium]]''.
* [[Caesalpinioideae]]: 148 genera and ~4400 species. [[Pantropical]]. ''[[Caesalpinia]]'', ''[[Senna (plant)|Senna]]'', ''[[Mimosa]]'', ''[[Acacia]]''. Includes the former subfamily [[Mimosoideae]] (80 genera and ~3200 species; mostly tropical and warm temperate Asia and America).
* [[Faboideae]] (Papilionoideae<ref>NOTE: The subfamilial name Papilionoideae for Faboideae is approved by the [[International Code of Nomenclature for algae, fungi, and plants]], Article 19.8</ref>): 503 genera and ~14,000 species. [[Cosmopolitan distribution|Cosmopolitan]]. ''[[Astragalus (plant)|Astragalus]]'', ''[[Lupinus]]'', ''[[Pisum]]''.