Icacinaceae

Icacinaceae
Icacina oliviformis
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Asterids
Order: Icacinales
Family: Icacinaceae
Miers[1]
Genera

See text.

The Icacinaceae are a family of flowering plants,[2] consisting of trees, shrubs, and lianas, primarily of the tropics.

The family was traditionally circumscribed quite broadly, with around 55 genera totalling over 400 species. In 2001, though, this circumscription was found to be polyphyletic, and the family was split into four families in three different orders: Icacinaceae sensu stricto (unplaced at order rank), Pennantiaceae (Apiales), Stemonuraceae (Aquifoliales) and Cardiopteridaceae (also Aquifoliales). Icacinaceae sensu stricto contains about 150 species, distributed into about 35 genera.[3] One genus was named in 2005.[4] Icacinaceae sensu stricto is not monophyletic,[5] but further studies will be needed before anyone can determine how it should be divided into segregate families. Icacina senegalensis extracts have shown activity against malaria parasites.[6]

Genera

The following list of genera was created from several sources. The last column contains those genera whose inclusion in the family is especially doubtful.

  • Alsodeiopsis
  • Casimirella
  • Chlamydocarya  
  • Desmostachys
  • Hosiea
  • Icacina
  • Iodes
  • Lavigeria
  • Leretia
  • Mappia
  • Mappianthus
  • Merrilliodendron  
  • Miquelia
  • Natsiatopsis
  • Natsiatum
  • Nothapodytes
  • Phytocrene
  • Pittosporopsis
  • Pleurisanthes
  • Polycephalium  
  • Polyporandra
  • Pyrenacantha
  • Rhyticaryum
  • Sarcostigma
  • Sleumeria

History

In 1841, George Bentham described Apodytes and Pogopetalum as new genera and united them with Icacina, Gomphandra, and Leretia to create the tribe Icacineae of what would later be called the family Olacaceae.[7] Olacaceae was at that time, and through the 20th century, defined broadly, encompassing several families in the order Santalales.[8] Pogopetalum was later synonymized with Emmotum.[9][10]

In 1852, John Miers argued that Bentham's Icacineae did not belong in Olacaceae and he raised them to the taxonomic rank of family as Icacinaceae.[11]

Philippe van Tieghem realized that the family Icacinaceae, as circumscribed by Miers, consisted of groups that were only distantly related, and in 1897, he divided it into seven families.[12][13] Van Tieghem's treatment in some ways anticipated the results of 21st century phylogenetic studies, in particular, by his establishment of the families Emmotaceae and Leptaulaceae. His division of Icacinaceae into smaller families was not accepted and other authors continued to define Icacinaceae in the broad sense, known as Icacinaceae sensu lato.

In 1942, Hermann Sleumer defined Icacinaceae broadly in his coverage of the family for Die Natürlichen Pflanzenfamilien.[14] Later authors did likewise.

In the 1940s, Richard A. Howard wrote a series of papers on several of the genera.[3] Sleumer wrote about the Asian genera in 1969,[15] and the Malesian genera in 1971.[16] Much of what is known about the family comes from regional floras such as Flora of Australia [17] and Flora of China.[18]

In 2001, Jesper Kårehed, using a combination of morphological and DNA sequence data, showed that Icacinaceae sensu lato was distantly polyphyletic and was, at least arguably, the worst of the plant families. It is now known to have rivaled Flacourtiaceae as an unnatural assemblage of disparate groups. Kårehed divided it into four families: Pennantiaceae, Stemonuraceae, Cardiopteridaceae, and Icacinaceae sensu stricto.[3]

Pennantiaceae consists of the single genus Pennantia and is the most basal clade in the campanulid order Apiales.[19][20]

Stemonuraceae is a family of 12 genera in the campanulid order Aquifoliales. It is sister to Cardiopteridaceae.[3][21]

Before the phylogeny produced by Kårehed, Cardiopteridaceae contained only Cardiopteris. Kårehed transferred Citronella, Gonocaryum, and Leptaulus from Icacinaceae to this family, and provisionally placed Metteniusa, Dendrobangia, and Pseudobotrys there as well. Metteniusa was shown to be a lamiid in 2007, and was placed in a family by itself.[22] The affinities of Dendrobangia and Pseudobotrys remain obscure.

Some authors have continued to maintain Cardiopteridaceae as a monogeneric family, placing Citronella, Gonocaryum, Leptaulus, Dendrobangia, and Pseudobotrys in Leptaulaceae.[23] The study by Kårehed showed Cardiopteris to be embedded in Leptaulaceae, but statistical support for this position was not strong.

Icacinaceae sensu stricto consists of about 35 genera of lamiids and is not a monophyletic group.[5] Icacina and some close relatives are probably related to the order Garryales and might well be placed within it. Cassinopsis, Emmotum, Apodytes, and a few others probably belong elsewhere among the lamiids. Apodytes and Rhaphiostylis might be closer to Oncothecaceae than to anything else.[5] The division of the lamiids into orders is still uncertain. More phylogenetic studies will be needed to resolve these issues.

References

  1. ↑ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III" (PDF). Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. Retrieved 2013-07-06.
  2. ↑ "Icacinaceae" At: Angiosperm Phylogeny Website At: Missouri Botanical Garden Website (see External links below).
  3. 1 2 3 4 Kårehed, Jesper (2001). "Multiple origin of the tropical forest tree family Icacinaceae". American Journal of Botany. 88 (12): 2259–2274. doi:10.2307/3558388. JSTOR 3558388. PMID 21669659.
  4. ↑ Timothy M.A. Utteridge, Hidetoshi Nagamasu, Stephen P. Teo, Lydia C. White, and Peter Gasson. 2005. "Sleumeria (Icacinaceae): A New Genus from Northern Borneo". Systematic Botany 30(3):635-643.
  5. 1 2 3 Frederick Lens, Jesper Kårehed, Pieter Baas, Steven Jansen, David Rabaey, Suzy Huysmans, Thomas Hamann, and Eric Smets. 2008. "The wood anatomy of the polyphyletic Icacinaceae s.l. and their relationships within asterids". Taxon 57(2):525-552.
  6. ↑ Sarr SO, Perrotey S, Fall I, Ennahar S, Zhao M, Diop YM, Candolfi E, Marchioni E.,"Icacina senegalensis (Icacinaceae), traditionally used for the treatment of malaria, inhibits in vitro Plasmodium falciparum growth without host cell toxicity." Malar J. 2011 Apr 11;10(1):85
  7. ↑ George Bentham. 1841. page 679. In: "Account of two new genera allied to Olacineae". Transactions of the Linnean Society of London 18:671-686 & plates 41 and 42. (see External links below).
  8. ↑ Valéry Malécot and Daniel L. Nickrent. 2008. "Molecular Phylogenetic Relationships of Olacaceae and Related Santalales". Systematic Botany 33(1):97-106.
  9. ↑ Rodrigo Duno de Stefano, Diego F. Angulo, and Fred W. Stauffer. 2007. "Emmotum harleyi, a New Species from Bahia, Brazil, and Lectotypification of Other Icacinaceae". Novon 17(3):306-309.
  10. ↑ Richard A. Howard. 1942. "Studies of the Icacinaceae. III. A revision of Emmotum". Journal of the Arnold Arboretum 23:479-494.
  11. ↑ John Miers. 1852. page 221. In: "Observations on the Affinities of the Icacinaceae". Annals and Magazine of Natural History, iccluding Zoology, Botany, and Geology, series 2. 9:218-226. (see External links below).
  12. ↑ Philippe E.L. van Tieghem. 1897. page 842. In: "Sur les inséminées à nucelle pourvu d'un seul tégument formant la subdivisions des Unitegminées ou Icacinées". Séance du Mardi 20 Avril 1897. Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences 124:839-844.
  13. ↑ Philippe E.L. van Tieghem. 1897. "Sur les phanerogams sans graines, formant la divisions des inséminées". Bulletin de la Société Botanique de France 44:99-139. (see External links below).
  14. ↑ Hermann Sleumer. 1942. "Icacinaceae" pages 322-396. In: H.G. Adolf Engler and Karl A.E. Prantl, with Hermann Harms and Johannes Mattfeld (editors). Die Natürlichen Pflanzenfamilien volume 20b. Duncker and Humblot: Berlin, Germany. 1960 reprint of 1942 publication.
  15. ↑ Hermann Sleumer. 1969. "Materials toward the knowledge of the Icacinaceae of Asia, Malesia, and adjacent areas". Blumea 17(1):181-264.
  16. ↑ Hermann Sleumer. 1971. "Icacinaceae" pages 1-87. In: Cornelis G.G.J. van Steenis (editor). Flora Malesiana series 1, volume 7. Noordhoff International Publishing: Leyden, Holland.
  17. ↑ Gordon P. Guymer. 1984. "Icacinaceae" pages 204-211. In: Alexander S. George (executive editor). Flora of Australia volume 22. Australian Government Publishing Service: Canberra, Australia.
  18. ↑ Hua Peng and Richard A. Howard. 2008. "Icacinaceae" pages 505-514. In: Zhengyi Wu, Peter H. Raven, and Deyuan Hong (editors). Flora of China volume 11. Science Press: Beijing, China; Missouri Botanical Garden Press: St. Louis, Missouri, USA.
  19. ↑ Jesper Kårehed. 2003. "The family Pennantiaceae and its relationships to Apiales". Botanical Journal of the Linnean Society 141(1):1-24.
  20. ↑ Gregory M. Plunkett, Gregory T. Chandler, Porter P. Lowry, Steven M. Pinney, and Taylor S. Sprenkle. 2004. "Recent advances in understanding Apiales and a revised classification". South African Journal of Botany 70(3):371-381.
  21. ↑ Douglas E. Soltis, Pamela S. Soltis, Peter K. Endress, and Mark W. Chase (2005). Phylogeny and Evolution of the Angiosperms. Sunderland, MA, USA: Sinauer. ISBN 978-0-87893-817-9.
  22. ↑ Favio González, Julio Betancur, Olivier Maurin, John V. Freudenstein, and Mark W. Chase. 2007. "Metteniusaceae, an early-diverging family in the lamiid clade". Taxon 56(3):795-800.
  23. ↑ Timothy M.A. Utteridge and Richard K. Brummitt. 2007. "Leptaulaceae" pages 191-192. In: Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. Flowering Plant Families of the World. Firefly Books: Ontario, Canada. (2007).
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