Cleomaceae

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids II
Ordo: Brassicales

Familia: Cleomaceae
Genera: Andinocleome – Areocleome – Arivela – Cleome – Cleomella – Cleoserrata – Coalisina – Cochranella – Corynandra – Dactylaena – Dipterygium – Gilgella – Gynandropsis – Haptocarpum – Iltisiella – Kersia – Melidiscus – Physostemon – Podandrogyne – Polanisia – Pterocleome – Puccionia – Rorida – Sieruela – Stylidocleome – Tarenaya – Thulinella
Name

Cleomaceae Horan.

Type genus: Cleome L. Sp. Pl. 2: 671–672. (1753) nom. cons.

Note: This is a taxonomic opinion summarised in Bayat et al. (2018), which is based upon a narrowly circumscribed Cleome s.s. However, an alternative view of Cleomaceae can be found on the Plants of the World site and in Christenhusz et al. (2018) this comprises only Cleome s.l., Cleomella and Dactylaena. The later circumscriptions finds many combinations unplaced and therefore can not be used here, as WS is a taxonomic database.
References
Primary references

Linnaeus, C. 1753. Species Plantarum. Tomus II: 671. Reference page.

Additional references

Barrett, R.L., Roalson, E.H., Ottewell, K., Byrne, M., Govindwar, S.P., Yadav, S.R., Tamboli, A.S. & Gholave, A.R. 2017. Resolving Generic Boundaries in Indian‐Australasian Cleomaceae: Circumscription of Areocleome, Arivela, and Corynandra as Distinct Genera. Systematic Botany 42(4): 694-708. DOI: 10.1600/036364417X696401 Reference page.
Bayat, S., Schranz, M.E., Roalson, E.H. & Hall, J.C. 2018. Lessons from Cleomaceae, the sister of crucifers. Trends in Plant Science 23(9): 808-821. DOI: 10.1016/j.tplants.2018.06.010 PDF Reference page.
Cardinal-McTeague, W.M., Sytsma, K.J. & Hall, J.C. 2016. Biogeography and diversification of Brassicales: a 103 million year tale. Molecular Phylogenetics and Evolution 99: 204-224. DOI: 10.1016/j.ympev.2016.02.021 PDF Reference page.
Christenhusz, M.J.M., Fay, M.F. & Byng, J.W. (eds). 2018. The Global Flora, Vol. 4: Special Edition, GLOVAP Nomenclature Part 1. Plant Gateway Ltd., Bradford. ISBN 978-0-9929993-5-3 PDFReference page.
Cochrane, T.S. & Iltis, H.H. 2014. Studies in the Cleomaceae VII: five new combinations in Corynandra, an earlier name for Arivela. Novon 23(1): 21–27. DOI: 10.3417/2013023 ResearchGate BHL Reference page.
Iltis, H.H. & Cochrane, T.S. 2007. Studies in the Cleomaceae V: A new genus and ten new combinations for the flora of North America. Novon 17(4): 447–451. DOI: 10.3417/1055-3177(2007)17[447:SITCVA]2.0.CO;2 BHL Reference page.
Iltis, H.H., Hall, J.C., Cochrane, T.S. & Sytsma, K.J. 2011. Studies in the Cleomaceae I: On the separate recognition of Capparaceae, Cleomaceae, and Brassicaceae. Annals of the Missouri Botanical Garden 98(1): 28–36. DOI: 10.3417/2007017 Reference page.
Iltis, H.H. & Cochrane, T.S. 2014. Studies in the Cleomaceae VI: a new genus and sixteen new combinations for the Flora Mesoamericana. Novon: A Journal for Botanical Nomenclature 23(1): 51-59. DOI: 10.3417/2013017 PDF from ResearchGate Reference page.
Inda, L.A., Torrecilla, P., Catalán, P. Ruiz-Zapata, T. 2008. Phylogeny of Cleome L. and its close relatives Podandrogyne Ducke and Polanisia Raf. (Cleomoideae, Cleomaceae) based on analysis of nuclear ITS sequences and morphology. Plant Systematics and Evolution 274(1-2): 111–126. DOI: 10.1007/s00606-008-0026-y Reference page.
Mcginty, E.M. & Roalson, E.H. 2020. Generic reorganization and nomenclatural synopsis of the Andean Clade (Cleomaceae). Phytotaxa 456(3): 256-268.DOI: 10.11646/phytotaxa.456.3.3 Paywall Reference page.
Patchell, M.J., Roalson, E.H. & Hall, J.C. 2014. Resolved phylogeny of Cleomaceae based on all three genomes. Taxon 63(2): 315–328. DOI: 10.12705/632.17 Full article (PDF) on ResearchGate Reference page.
Roalson, E.H. 2021. A revised synonymy, typification, and key to species of Cleome sensu stricto (Cleomaceae). Phytotaxa 496(1): 54-68. DOI: 10.11646/phytotaxa.496.1.2 Paywall Reference page.
Roalson, E.H., Hall, J.C., Riser, J.P., Cardinal-McTeague, W.M., Cochrane, T.S. & Sytsma, K.J. 2015. A revision of generic boundaries and nomenclature in the North American cleomoid clade (Cleomaceae). Phytotaxa 205(3): 129–144. DOI: 10.11646/phytotaxa.205.3.1 ResearchGate Reference page.
Roalson, E.H. & Hall, J.C. 2017. New generic concepts for African Cleomaceae. Systematic Botany 42(4): 925–942. DOI: 10.1600/036364417X696393 BioOne ResearchGate Reference page.
Roalson, E.H. & Neto, R.L.S. 2020. Nomenclatural notes in Physostemon (Cleomaceae). Phytotaxa 468(3): 275-282. DOI: 10.11646/phytotaxa.468.3.3 Paywall Reference page.
Thulin, M. & Roalson, E.H. 2017. Resurrection of the genus Rorida (Cleomaceae), a distinctive Old World segregate of Cleome. Systematic Botany 42(3): 569–577. DOI: 0.1600/036364417X695989 ResearchGate Reference page.
Tucker, G.C., Vanderpool, S.S. 2010. Cleomaceae. Pp. 199–223 in: Flora of North America Editorial Committee (eds.), Flora of North America north of Mexico, vol. 7, Magnoliophyta: Salicaceae to Brassicaceae. New York: Oxford University Press. [Not seen]

Links

Govaerts, R. et al. 2019. Cleomaceae in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2019 June 8. Reference page.
Hassler, M. 2019. Cleomaceae. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2019. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: 2019 June 4. Reference page.
International Plant Names Index. 2021. Cleomaceae. Published online. Accessed: Sept 14 2021.
Tropicos.org 2014. Cleomaceae. Missouri Botanical Garden. Published online. Accessed: 14 Sept. 2014.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.

Vernacular names
العربية: ذفرية
català: Cleomàcies
čeština: Luštěnicovité
suomi: Hämähäkkikukkakasvit
français: Cléomacées
日本語: フウチョウソウ科
한국어: 풍접초과
русский: Клеомовые
svenska: Paradisblomsterväxter
ไทย: วงศ์ผักเสี้ยน
Tiếng Việt: Họ Màn màn
中文: 醉蝶花科, 白花菜科

The Cleomaceae are a small family of flowering plants in the order Brassicales, comprising about 300 species in 10 genera, or about 150 species in 17 genera.[2] These genera were previously included in the family Capparaceae, but were raised to a distinct family when DNA evidence suggested the genera included in it are more closely related to the Brassicaceae than they are to the Capparaceae. The APG II system allows for Cleomaceae to be included in Brassicaceae.[3]

Taxonomy

In 1994, a group of scientists including Walter Stephen Judd suggested to merge the Capparaceae (which at that time included the Cleomaceae) with the Brassicaceae. Early DNA-analysis showed that the Capparaceae - as defined at that moment - were paraphyletic, and others suggested to assign the genera closest to the Brassicaceae to the Cleomaceae.[4] The Cleomaceae and Brassicaceae diverged approximately 41 million years ago.[5] All three families have consistently been placed in one order (variably called Capparales or Brassicales).[4] The APG II system merged Cleomaceae and Brassicaceae. Other classifications have continued to recognise the Capparaceae, but with a more restricted circumscription, either including Cleome and its relatives in the Brassicaceae or recognizing them in the segregate family Cleomaceae. The APG III system has recently adopted this last solution, but this may change as a consensus arises on this point. Current insights in the relationships of the Brassicaceae, based on a 2012 DNA-analysis, are summarized in the following tree.[6]

core Brassicales	family Resedaceae       family Gyrostemonaceae       family Pentadiplandraceae         family Tovariaceae       family Capparaceae       family Cleomaceae     family Brassicaceae             family Emblingiaceae

 
Subdivision

The genera in Cleomaceae include:[7]

Andinocleome
Arivela
Buhsea
Carsonia
Chilocalyx
Cleome
Cleomella
Cleoserrata
Dactylaena
Decastemon
Gynandropsis
Haptocarpum
Hemiscola
Iltisiella
Isomeris
Justago
Melidiscus
Mitostylis
Oxystylis
Peritoma
Physostemon
Podandrogyne
Polanisia
Rorida
Roridula Forssk.
Tarenaya
Tetratelia
Wislizenia

Description
Floral diagram of Cleome hassleriana

Cleomaceae are mostly annual, sometimes perennial herbaceous plants, seldom shrubs or small trees, and some are lianas. Most of them have glands, a character setting them apart from the related Capparaceae, and contain resins, and may have an aromatic or foetid smell. The alternately set leaves, have stalks and never form a sheath around the stem. The leaves are always palmately compound, mostly with three to seven leaflets, rarely with just one leaflet. At the base of the leafstalk sit scaly, leafy or spiny stipules, a difference with its sister group Brassicaceae that consistently lack stipules, although a pair of glands can sometimes be present at the base of the leaf stalk. Hairs and papillae may be present.[8]

The flowers of Cleomaceae are somewhat to strongly bilateral symmetric. There are mostly four, sometimes two or six, free or merged sepals, that may be shed quickly or remain, are overlapping in the bud or are already separate. The corolla consists of mostly four, sometimes two or six petals that have distinctive claws and limbs, free or with merged limbs. The stamens may be placed on the level of insertion of the perianth or on a stalk, raising the bases of the stamens above the perianth. This occurs for instance in Gynandropsis gynandra. There are four simple stamens or branched in four groups of up to 12 stamens each that have a common base, and are free from the petals. When there are groups of stamens, the anthers mature from the middle outwards. The base of the stamens may or may not be fused with the female parts in the centre of the flower. The filaments may be of equal length or strongly differ, and all may carry anthers, but infertile staminodes may also occur. The anther releases the pollen through slits stretching along its length. The ovary is positioned above the perianth and the androecium (a so-called superior ovary), consists of two fully merged carpels, and contains one cavity that may be divided in two by a partition (or replum). Between ten and fifty ovules (future seeds) are set along the margins of each of the carpels. As the fruit ripens, a stalk (or gynophore) develops. The dry fruit opens with two valves. When the fruit has a replum, it has the same basic morphology as the Brassicaceae (called silicula or siliqua), and this sets it apart from the Capparaceae, that have a capsule.[8]
References

Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
Cleomaceae, Zhang Mingli (张明理)1; Gordon C. Tucker2, Harvard.edu, [1]
Stevens, P. F. (2001 onwards) Angiosperm Phylogeny Website: Brassicales.
Hall, J.C.; Sytsma, K.J.; Iltis, H.H. (2002). "Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data". American Journal of Botany. 89 (11): 1826–1842. doi:10.3732/ajb.89.11.1826. PMID 21665611.
Renate Schmidt; Ian Bancroft, eds. (2010). Genetics and Genomics of the Brassicaceae. Plant Genetics and Genomics: Crops and Models. Vol. 9. Springer Science & Business Media.
Su, Jun-Xia; Wang, Wei; Zhang, Li-Bing; Chen, Zhi-Duan (June 2012). "Phylogenetic placement of two enigmatic genera, Borthwickia and Stixis, based on molecular and pollen data, and the description of a new family of Brassicales, Borthwickiaceae" (PDF). Taxon. 61 (3): 601–611. doi:10.1002/tax.613009. Archived from the original (PDF) on 2017-08-03. Retrieved 2017-08-17.
Germplasm Resources Information Network: Cleomaceae
Watson, L.; Dallwitz, M.J. (1992). "Cleomaceae (Pax) Airy Shaw". The families of flowering plants. Retrieved 2018-10-11.