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Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Ordo: Santalales

Familia: Balanophoraceae
Genera: Balanophora – Chlamydophytum – Corynaea – Dactylanthus – Ditepalanthus – Exorhopala – Hachettea – Helosis – Langsdorffia – Lathrophytum – Lophophytum – Mystropetalon – Ombrophytum – Rhopalocnemis – Sarcophyte – Scybalium – Thonningia
Name

Balanophoraceae L.C. & A. Rich. Mém. Mus. Hist. Nat. 8: 429. (1822) nom. cons.

Type genus: Balanophora J.R.Forst. & G.Forst. Char. Gen. Pl. 50. (1775)

Synonyms

Heterotypic
Mystropetalaceae (Engl.) Takht., Bot. Zhurn. (Moscow & Leningrad) 75(5): 698. (1990)

Note: Su et al. (2015) prefer to segregate Dactylanthus, Hachettea and Mystropetalon into Mystropetalaceae, but this has not gained wider acceptance and so a conservative position is adopted here.
References

Richard, L.C.M. 1822. Mémoires du Muséum d'Histoire Naturelle 8: 429.
Nickrent, D.L., Soltis, D.E., Colwell, E.A., Wolfe, A.D., Young, N.D., Steiner, K.E., & de Pamphilis, C.W. 1998. Molecular phylogenetic and evolutionary studies of parasitic plants. Pp. 211-241, in Soltis, D. E., Soltis, P. S., & Doyle, J. J. (eds), Molecular Systematics of Plants II: DNA Sequencing. Kluwer, Boston.
Nickrent, D.L., Malécot, V., Vidal-Russell, R. & Der, J.P. 2010. A revised classification of Santalales. Taxon 59(2): 538–558. DOI: 10.1002/tax.592019 JSTOR PDF. Reference page.
Nickrent, D.L. 1997-2019. The Parasitic Plant Connection, Department of Plant Biology, Southern Illinois University Carbondale. Online. Reference page.
Hassler, M. 2019. Balanophoraceae. 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 on the internet. Accessed: 2019 Jan. 25. Reference page.
Lohmüller, F.A. 2005. The Botanical System of the Plants[1]
Su, H.J., Hu, J.M., Anderson, F.E., Der, J.P. & Nickrent, D.L. 2015. Phylogenetic relationships of Santalales with insights into the origins of holoparasitic Balanophoraceae. Taxon 64(3): 491-506. DOI: 10.12705/643.2 PDF from ResearchGate Reference page.

Vernacular names
日本語: ツチトリモチ科
中文: 蛇菰科

The Balanophoraceae are a subtropical to tropical family of obligate parasitic flowering plants, notable for their unusual development and formerly obscure affinities. In the broadest circumscription, the family consists of 16 genera. Alternatively, three genera may be split off into the segregate family Mystropetalaceae.

The plants are normally found in moist inland forests growing on tree roots and have an above ground inflorescence with the overall appearance of a fungus, composed of numerous minute flowers. The inflorescences develop inside the tuberous underground part of the plant, before rupturing it and surfacing. The plants are monoecious, or dioecious, and the fruits are indehiscent drupes or nuts. The underground portion, which attaches itself to the host, looks like a tuber, and is not a proper root system. The plants contain no chlorophyll. Balanophora means "bearing an acorn" (from the shape of the female inflorescence).

Taxonomy

In the classification system of Dahlgren, the Balanophoraceae were placed in the order Balanophorales in the superorder Balanophoriflorae (also called Balanophoranae). The APG IV system of 2016 (unchanged from the APG III system of 2009[2]), also recognizes this family, including it in the order Santalales,[3] where it was also placed by the Cronquist system (1981).

A 2015 molecular phylogenetic suggested that as circumscribed in the APG IV system, Balanophoraceae is not monophyletic. The authors of the study proposed dividing Balanophoraceae s.l. into Balanophoraceae s.s. and the family Mystropetalaceae, containing three monotypic genera, Dactylanthus, Hachettea and Mystropetalon. The cladogram below shows the relationships obtained (using the broad APG IV circumscriptions of Olacaceae and Santalaceae).[4][5]
Santalales

Olacaceae s.l.


Balanophoraceae s.s.




Mystropetalaceae

Loranthaceae



Misodendraceae

Schoepfiaceae




Opiliaceae

Santalaceae s.l.






Balanophoraceae s.l.
Genera

As of July 2021, the broadly circumscribed family was accepted by Plants of the World Online, which included the following genera.[1] Other sources place three of the genera in the segregate family Mystropetalaceae.[4][5]

Balanophora J.R.Forst. & G.Forst.
Chlamydophytum Mildbr.
Corynaea Hook.f.
Dactylanthus Hook.f. [Mystropetalaceae]
Ditepalanthus Fagerl.
Hachettea Baill. [Mystropetalaceae]
Helosis Rich.
Langsdorffia Mart.
Lathrophytum Eichler
Lophophytum Schott & Endl.
Mystropetalon Harv. [Mystropetalaceae]
Ombrophytum Poepp. ex Endl.
Rhopalocnemis Jungh.
Sarcophyte Sparrm.
Scybalium Schott & Endl.
Thonningia Vahl

References

"Balanophoraceae Rich.". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2021-07-20.
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.
Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.
Su, Huei-Jiun; Hu, Jer-Ming; Anderson, Frank E.; Der, Joshua P. & Nickrent, Daniel L. (2015). "Phylogenetic relationships of Santalales with insights into the origins of holoparasitic Balanophoraceae" (PDF). Taxon. 64 (3): 491–506. doi:10.12705/643.2. Retrieved 2021-07-20.
Nickrent, Daniel L. (2020). "Parasitic angiosperms: How often and how many?". Taxon. 69 (1): 5–27, S3. doi:10.1002/tax.12195.

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