Celastrales
| Celastrales | |
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| Euonymus europaea, Celastraceae family | |
| Scientific classification | |
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| Order: | Celastrales Baskerville
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| Families | |
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Celastraceae (staff vine family) | |
Celastrales is an order of flowering plants. They are found throughout the tropics and subtropics, with only a few species extending far into the temperate regions. There are about 1200 [1] to 1350 [2] species in about 100 genera. All but 7 of these genera are in the large family Celastraceae. Until recently, the composition of the order and its division into families varied greatly from one author to another.
Description
Celastrales is a diverse order that has no conspicuous unique characteristic and is consequently hard to recognize. [3] The flowers are usually small with a conspicuous nectary disk. The stipules are small or rarely absent. The micropyle has 2 openings and is therefore called a bistomal micropyle. Flowers with well-developed male and female parts are often functionally unisexual. The seed often has an aril. In bud, the sepals have a quincuncial arrangement. This means that 2 sepals are inside, 2 are outside, and the other has one side covered and one side exposed.
Relationships
Perhaps the most conspicuous and unusual trait of Celastrales is the nectary disk, a feature that it shares with another rosid order, Sapindales. Since the 2 orders are not closely related, the disk must have been an independent development in each of these 2 lines.
Celastrales is a member of the COM clade [4] of Fabidae, with Fabidae being one of the 2 groups of Eurosids. [5] The COM clade consists of 3 orders, Celastrales, Oxalidales, Malpighiales, and 1 unplaced family, Huaceae. Hardly anything is known about how the 4 parts of the COM clade are related to each other.
Circumscription
The name Celastraceae was first used by Thomas Baskerville in 1839. [6] In the time since Baskerville first defined the order, until the 21st century, there were great differences of opinion about what should be included in the order and in its largest family, Celastraceae. The family Celastraceae was the only group consistently placed in the order by all authors. Because of the ambiguity and complexity of its definition, Celastraceae became a dumping ground for genera of dubious affinity. Several genera were assigned to this family with considerable doubt about whether they really belonged there. Also, some genera that properly belong in Celastraceae were placed elsewhere.
By the end of the 20th century, Goupia and Forsellesia had been excluded from Celastraceae and also from Celastrales. Goupia is now in Malpighiales. [7] Forsellesia is now in Crossosomatales. [8] It continues to be the subject of a dispute about whether its proper name is Forsellesia or Glossopetalon. [9]
After being placed elsewhere, Canotia, Brexia, and Plagiopteron were found to belong in Celastraceae. The family Hippocrateaceae was found to be deeply nested within Celastraceae and is no longer recognized as a separate family.
In 2000, Vincent Savolainen and co-authors found that 3 families - Lepidobotryaceae, Parnassiaceae, and Celastraceae - were closely related and they stated that these should constitute the order Celastrales. [10] They also excluded Lophopyxis from Celastrales. Lophopyxis now constitutes a monogeneric family in Malpighiales. [7] In a DNA study in 2001, Mark Simmons and others confirmed all of these results except for the placement of Lophopyxis and Lepidobotryaceae, which they did not sample. [11]
In 2006, Li-Bing Zhang and Mark Simmons produced a phylogeny of Celastrales based on nuclear ribosomal and chloroplast DNA. [12] Their results showed that Bhesa and Perrottetia were misplaced in Celastraceae. Bhesa is now in Malpighiales. [7] and Perrottetia is in Huerteales. [13] Zhang and Simmons found Pottingeria and Mortonia to be closely related to the families Parnassiaceae and Celastraceae, but not in either of them. These two genera are therefore in Celastrales. They found that Siphonodon and Empleuridium are proper members of Celastraceae, removing considerable doubt about their placement there. They also showed that the small family Stackhousiaceae, consisting of three genera, is embedded in Celastraceae. Most of these results were confirmed by the second phylogeny of Celastrales, which was produced by Mark Simmons and several co-authors in 2008. [14]
Nicobariodendron sleumeri, the only member of its genus, continues to be an enigma. It is a small tree from the Andaman and Nicobar Islands of India. Little is known of it and it has never been sampled for DNA. It is generally thought to belong in Celastrales, [2] but this is not a certainty.
Families
Celastrales has been divided into families in various ways. In their APG II classification in 2003, the Angiosperm Phylogeny Group recognized 3 families in Celastrales - Lepidobotryaceae, Parnassiaceae, and Celastraceae. At the Angiosperm Phylogeny Website, Peter F. Stevens has recognized these 3 families and added a 4th, Pottingeriaceae, consisting only of Pottingeria.
Lepidobotryaceae has 2 species, Lepidobotrys staudtii, and Ruptiliocarpon caracolito. Parnassiaceae has 2 genera. Lepuropetalon is monospecific, (Lepuropetalon spathulatum) and Parnassia has about 70 species.
In the 2006 phylogeny, Nicobariodendron was not sampled, but those species that were sampled fell into 2 strongly supported clades. One was a small clade consisting only of the family Lepidobotryaceae. The other was a very large clade containing the rest of the order. They found the large clade to be divided into 5 strongly supported groups. These are the family Parnassiaceae, the genus Pottingeria, the genus Mortonia, a pair of genera from Celastraceae (Quetzalia and Zinowiewia), and the rest of Celastraceae. No relationships were resolved among these 5 groups.
In 2008, Simmons and others produced a phylogeny of Celastrales that achieved better resolution than the 2006 study by sampling more species and more DNA. In the large clade of Celastrales, they found the same 5 groups that were found in the 2006 study. However, there was weak to moderate support for a clade of (Pottingeria + Mortonia), and weak support (56% maximum likelihood bootstrap percentage) for a clade containing Quetzalia, Zinowiewia, and the rest of Celastrales.
The division of Celastrales into families is clearly in need of revision, but further studies will be needed to determine the best way to do so. It is not yet clear whether Pottingeria and Mortonia are best placed into monogeneric families or combined into a single family. The weak support for the inclusion of Quetzalia and Zinowiewia in Celastraceae is a bit of a surprise, since neither had previously been thought to be an anomalous member of the family. An alternative approach would be to expand Celastraceae even further, to include Mortonia, Pottingeria, and Parnassiaceae.
Phylogeny
The following phylogenetic tree was made by combining parts of three different trees. [11] [12] [14] Bootstrap support is 100% except where shown. Branches with less than 50% bootstrap support are collapsed. The clade numbers are from Simmons et alii (2008).
| Lepidobotryaceae | ||||||||||||||||||||||||||||||||||
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References
- ^ "Lepidobotryaceae", "Parnassiaceae", and "Celastraceae" In: Klaus Kubitzki (ed.). The Families and Genera of Vascular Plants vol. VI. Springer-Verlag: Berlin,Heidelberg (2004).
- ^ a b Peter F. Stevens (2001 onwards). Angiosperm Phylogeny Website. In: Missouri Botanical Garden Website
- ^ Merran L. Matthews and Peter K. Endress (2005). "Comparative floral structure and systematics in Celastrales". Botanical Journal of the Linnean Society 149 (2):129-194
- ^ Hengchang Wang, Michael J. Moore, Pamela S. Soltis, Charles D. Bell, Samuel F. Brockington, Roolse Alexandre, Charles C. Davis, Maribeth Latvis, Steven R. Manchester, and Douglas E. Soltis (2009). "Rosid radiation and the rapid rise of angiosperm-dominated forests". Proceedings of the National Academy of Sciences 106(10):3853-3858. 10Mar2009.
- ^ Philip D. Cantino, James A. Doyle, Sean W. Graham, Walter S. Judd, Richard G. Olmstead, Douglas E. Soltis, Pamela S. Soltis, and Michael J. Donoghue (2007). "Towards a phylogenetic nomenclature of Tracheophyta". Taxon 56(3):822-846.
- ^ Thomas Baskerville. Affinities of Plants: with some observations upon progressive development. Taylor and Walton: Gower Street, London. (1839).
- ^ a b c Kenneth J. Wurdack and Charles C. Davis (2009). "Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm tree of life".
- ^ Robert F. Thorne and Ron Scogin (1978). Forsellesia Greene (Glossopetalon Gray), a third genus in the Crossosomataceae (Rosinae, Rosales). Aliso 9(2):171-178.
- ^ Victoria Sosa. "Crossosomataceae" In: Klaus Kubitzki (ed.) The Families and Genera of Vascular Plants vol.IX. Springer-Verlag: Berlin,Heidelberg (2007).
- ^ Vincent Savolainen, Michael F. Fay, Dirk C. Albach, Anders Backlund, Michelle van der Bank, Kenneth M. Cameron, S.A. Johnson, M. Dolores Lledo, Jean-Christophe Pintaud, Martyn P. Powell, Mary Clare Sheahan, Douglas E. Soltis, Pamela S. Soltis, Peter Weston, W. Mark Whitten, Kenneth J. Wurdack and Mark W. Chase (2000). "Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences". Kew Bulletin 55(2):257-309.
- ^ a b Mark P. Simmons, Vincent Savolainen, Curtis C. Clevinger, Robert H. Archer, and Jerrold I. Davis (2001). "Phylogeny of Celastraceae Inferred from 26S Nuclear Ribosomal DNA, Phytochrome B, rbcL, atpB, and Morphology". Molecular Phylogenetics and Evolution 19(3):353-366. doi:10.1006/mpev.2001.0937
- ^ a b Li-Bing Zhang and Mark P. Simmons (2006). "Phylogeny and Delimitation of the Celastrales Inferred from Nuclear and Plastid Genes". Systematic Botany 31(1):122-137.
- ^ Andreas Worberg, Mac H. Alford, Dietmar Quandt, and Thomas Borsch (2009). Huerteales sister to Brassicales plus Malvales, and newly circumscribed to include Dipentodon, Gerrardina, Huertea, Perrottetia, and Tapiscia. Taxon 58(in press).
- ^ a b Mark P. Simmons, Jennifer J. Cappa, Robert H. Archer, Andrew J. Ford, Dedra Eichstedt, and Curtis C. Clevinger (2008). "Phylogeny of the Celastreae and the relationships of Catha edulis inferred from morphological characters and nuclear and plastid genes". Molecular Phylogenetics and Evolution 48(2):745-757.
External links
- Phylogeny 2008 Mark P. Simmons, et alii (2008). Mol.Phyl.Evol. 48(2):745-757.
doi:10.1016/j.ympev.2008.04.039