Ira B. Glassman
BIO 499
Professor Clark
December 8th, 2016
1 Q: Family name, with author citation (from IPNI)
A: Fabaceae Lindl., 1836.[1] (Leguminosae Jussieu, nom. cons.).[2]
2 Q: Type genus, and ordinarily the type species (if you’re having trouble hacking through a nomenclatural jungle to get the type species, contact me well in advance of the due date)
A: Type genus: Faba Mill. (1754), nom. illeg. [= Vicia L. 1753).[1]
A: Type Species: Vicia faba L.[3]
3 Q: Alternate names for the family that include the type genus (homotypic synonyms). Most often, there aren’t any.
A: Leguminosae Jussieu, nom. cons.[2]
4 Q: The position of the family in each of these classifications (i.e., its order, class, and division/phylum), or if it doesn’t occur, an explanation of why.
A: Bessey:
Phylum – Anthophyta[4]
Class – Oppositifoliae (Dicotyledoneae)[4]
Subclass — Oppositifoliae-Cotyloideae[4]
Superorder — Cotyloideae-Apopetalae[4]
Order – Rosales[4]
A: Takhtajan:
Division – Magnoliophyta or Angiospermae[5]
Class – Magnoliopsida or Dicotyledones[5]
Subclass – Rosidae[5]
Superorder — Rosanae[5]
Order – Fabales[5]
Suborder – Saxifragineae[5]
A: Thorne:
Class – Angiospermae (Annonopsida)[6]
Subclass – Dicotyledoneae (Annonidae)[6]
Superorder – Rosineae[6]
Order – Rosales[6]
Suborder – Rosineae[6]
A: Dahlgren:
Class – Magnoliopsida (=Angiospermae)[7]
Subclass – Magnoliidae (=Dicotyledoneae)[7]
Superorder – Fabiflorae[7]
Order – Fabales[7]
A: APG I:
Clade – Eurosids I[8]
Order – Fabales Bromhead[8].
Only the family names, Orders, and clades are present within APG I. The classification believes that the family is, “central in flowering plant systematics.”[8] Instead of including Class, Phylum, or Divisions, APG I uses a more broad circumscription of the orders. [8]
A: APG IV:
Clade – Rosids[9]
Order – Fabales Bromhead.[9]
APG IV is the same as APG I in that it only lists Clades, Orders, and Families.
5 Q: Discussion of changes in circumscription (the genera that are included along with the type genus). Some families have retained the same circumscription for a century or more. Others are either more inclusive or less inclusive in APG as a result of maintaining monophyly. For large families, you obviously don’t need to include all the genera, but you should pick enough to show trends.
A: In Bessey’s 1915 classification, Fabaceae contained no subfamilies and was composed of the type genus Vicia, as well as a few other genera like Astragalus and Pisum.[4] In 1976, Thorne composed a classification where Fabaceae contained three subfamilies: Caesalpinoideae, Mimosoideae, and Faboideae.[6] In 1980, Dahlgren proposed a classification system which separated the families Mimosaceae, Caesalpiniaceae, and Fabaceae.[7] In APG III, the three subfamilies from Thorne’s classification returned to Fabaceae, where the genera in Vicia, Astragalus, Pisum and other genera from Bessey’s classification were included in Faboideae.[2] Mimosoideae and Caesalpiniodeae were composed of other genera added to the family like Mimosa and Caesalpinia.[2] Modern molecular phylogenetics have shown that the Faboideae and Mimososoideae stem from Ceasalpinioideae, composing it’s own clade, Caesalpinieae.[14]
6 Q: Related to the previous point, what other families have been split from your family, and what have been combined with your family, in different classifications.
A: The family Fabaceae, according to APG III, are comprised of three subfamilies: Mimosoideae (80 genera), Caesalpiniodeae (170 genera), and Faboideae (470 genera).[2] In Cronquist and Dahlgren classification systems, the three subfamilies rose to the family level, leaving a less inclusive Fabaceae family.[2]
The remainder of the information will come from APG IV and the references it cites (including previous APG publications and other papers in the primary literature).
7 Q: What is the phylogenetic outgroup to your family? It may be another family, an order, an unranked clade, or even a genus. What is the evidence (direct synapomorphy, or gene trees) that links your family to its outgroup?
A: A phylogenetic outgroup to Fabaceae is Quillajaceae.[2] Fabaceae and Quillajaceae are linked together by a unique trait in the order Fabales. A single carpel develops into a fruit, which dehisces along one or two edges at maturity, expelling developed seeds from the ovules.[10]
8 Q: What are the other families in the same order as your family? There may be none, and there may still be families that are not assigned to an order. Are there any phenotypic synapomorphies for the order?
A: The other families within the order Fabales are: Quillajaceae, Surianaceae, and Polygalaceae.[9] A phenotypic synapomorphy of the order Fabales is a part of the Nitrogen Fixing clade of plants, which develop root nodules with Rhizobium, which help to maintain nitrogen concentration within soil.[10][11] The order can also be classified with features: vessel elements with simple perforation plates, wood that is often fluorescing, and they also produce chlorophyllous embyros.[12]
9 Q: What are the phenotypic synapomorphies for your family? Are there any members in which any of these synapomorphies is not apparent? Why?
A: The plants possess intermediate inflorescences, which are occasionally reduced to a single flower.[2] The flowers contain a short hypanthium and a single carpel.[2] Fertilization produces fruits called legumes.[2] All of the Fabaceae subfamilies form root nodules for nitrogen fixation, however it is far less common in Caesalpinioideae.[2] The family also produces chemical substances for predator defense.[10] The production of alkaloids, flavonoids, tannins and free amino acid canavanine are commonly used in legume classification.[10]
10 Q: Following up on item 6 above, for each of the subdividions of your family (e.g., subfamilies) that were considered to be separate families in earlier classifications, explain whether they are monophyletic, paraphyletic, or polyphyletic, and (especially for the monophyletic ones) why APG chose to combine them.
A: Mimosoideae and Faboideae are monophyletic, the Caesalpinioideae are paraphyletic.[2] Results from plastid rbcL gene analysis confirmed that Mimosoideae and Faboideae are monophyletic subfamilies nested within a paraphyletic Caesalpinioideae.[13] The paraphyletic Caesalpinioideae compose a clade, earlier referred to in question 5 as Caesalpinieae. Due to the circumscription remaining the same throughout APG publications, there is no mention of why APG chose to combine them.
11 Q: Do you think the current circumscription of the family is adequate, or is it likely to change with more data?
A: Throughout the late 20th and early 21st century, studies stemming from global research utilizing morphology, DNA data, and cladistic analysis have been used to confirm the current circumscription of the family.[2] These studies have shown that Mimosoideae and Faboideae are both monophyletic, nested within a paraphyletic sub-family Caesalpinioideae. The agreement to keep Mimosoideae, Caesalpinioideae, and Faboideae at the sub-family level is not likely to change.[2]
References
1. http://ipni.org/ipni/idPlantNameSearch.do?id=30000147-2&back_page=%2Fipni%2FeditAdvPlantNameSearch.do%3Ffind_infragenus%3D%26find_isAPNIRecord%3Dtrue%26find_geoUnit%3D%26find_includePublicationAuthors%3Dtrue%26find_addedSince%3D%26find_family%3DFabaceae%26find_genus%3D%26find_sortByFamily%3Dtrue%26find_isGCIRecord%3Dtrue%26find_infrafamily%3D%26find_rankToReturn%3Dall%26find_publicationTitle%3D%26find_authorAbbrev%3D%26find_infraspecies%3D%26find_includeBasionymAuthors%3Dtrue%26find_modifiedSince%3D%26find_isIKRecord%3Dtrue%26find_species%3D%26output_format%3Dnormal.
2. https://en.wikipedia.org/wiki/Fabaceae
3. http://ipni.org/ipni/idPlantNameSearch.do?id=30327915-2&back_page=%2Fipni%2FeditAdvPlantNameSearch.do%3Ffind_infragenus%3D%26find_isAPNIRecord%3Dtrue%26find_geoUnit%3D%26find_includePublicationAuthors%3Dtrue%26find_addedSince%3D%26find_family%3D%26find_genus%3DFaba%26find_sortByFamily%3Dtrue%26find_isGCIRecord%3Dtrue%26find_infrafamily%3D%26find_rankToReturn%3Dall%26find_publicationTitle%3D%26find_authorAbbrev%3D%26find_infraspecies%3D%26find_includeBasionymAuthors%3Dtrue%26find_modifiedSince%3D%26find_isIKRecord%3Dtrue%26find_species%3D%26output_format%3Dnormal
4. Charles E. Bessey
Annals of the Missouri Botanical Garden Vol. 2, No. 1/2, Anniversary Proceedings (Feb. – Apr., 1915), pp. 109-164
5. Armen L. Takhtajan
Botanical Review Vol. 46, No. 3, Outline of the Classification of Flowering Plants (Magnoliophyta) (Jul. – Sep., 1980), pp. 225-359
6. Robert F. Thorne
Evolutionary Biology Vol. 9, A Phylogenetic Classification of the Angiospermae (1976), pp. 35-106
7. DAHLGREN, R. M. T. (1980), A Revised System of Classification of
the Angiosperms. Botanical Journal of the Linnean Society, 80: 91–124. doi:10.1111/j.1095-8339.1980.tb01661.x
8. The Angiosperm Phylogeny Group Annals of the Missouri Botanical
Garden Vol. 85, No. 4 (1998), pp. 531-553
9. The Linnean Society of London, Botanical Journal of the
Linnean Society, 2016, 181, 1–20
10. https://www.britannica.com/plant/Fabales
11. http://angio.bergianska.se/rosids/Polygalales/Polygalales.html
12. http://www.mobot.org/MOBOT/research/APweb/
13. http://tolweb.org/Fabaceae/21093/2006.06.14
14. https://en.wikipedia.org/wiki/Caesalpinioideae