Lilioid monocots
Lilioid monocots | |
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Lilium candidum | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Monocots |
Grade: | Lilioid monocots |
Orders | |
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Lilioid monocots (lilioids, liliid monocots, petaloid monocots, petaloid lilioid monocots) is an informal name used for a
The development of
Description
True lilioids
The descriptive term "petaloid lilioid monocot" relates to the conspicuous petal-like (
However, floral
Various trends are apparent among the lilioids, notably a change to an
Comparison with other monocot orders
In the orders that branched off before the lilioid monocots, the
The later evolved commelinids have various kinds of flower, few of which are 'lily-like'. In the order Poales, comprising grasses, rushes and sedges, flowers are either petal-less or have small, unshowy petals. Many Zingiberales species have brightly coloured and showy flowers. However, their apparent structure is misleading. For example, the six tepals of cannas are small and hidden under expanded and brightly coloured stamens or staminodes which resemble petals and may be mistaken for them.[8]
Taxonomy
Early history
In one of the earliest monocot taxonomies, that of
The term "lilioid monocot" or lilioid" has had widely varying interpretations.
As Kron and
The development of
Phylogenetic era
In the 1995 study by Chase et al. referred to above, which was the largest yet to use purely molecular data, the results demonstrated paraphyly of the lilioids. However, because their data contradicted purely morphological phylogenies they were reluctant to draw definite conclusions as to the monophyly of this group.
There was no clear clade corresponding to Dahlgren's Liliiflorae, whose families were distributed amongst the aroids and dioscoreoids. Of Dahlgren's Liliiflorae, the Dioscoreales largely grouped into dioscoreoids, with the exception of Stemonaceae. The Asparagales formed two major groupings, which they labelled "higher" and "lower asparagoids", and included both the Iridaceae and Orchidaceae from Dahlgren's Liliales. On the other hand, a number of families from three other orders (Asparagales, Dioscoreales, Melanthiales) segregated together with the remaining Liliales families. Genera from Dahlgren's Melanthiales were found in both dioscoreoids and the redefined Liliales. Finally Dahlgren's Burmanniales were found to belong with the dioscoreoids. Some Asparagales taxa were also found amongst the commelinoids. The stemonoids were formed from Stemonaceae and other families from a variety of orders, including Pandanaceae (which alone formed Dahlgren's Pandaniflorae).
In an attempt to resolve the apparent differences between morphological and molecularly defined trees, a combined analysis was undertaken[c] which confirmed superorder Liliiflorae as monophyletic, provided that a few modifications were undertaken. These included the removal of two tribes of Melanthiaceae (Melanthiales) and the inclusion of three additional families (Cyclanthaceae, Pandanaceae and Velloziaceae) from other superorders. This newly and more narrowly redefined Lilianae/Liliiflorae contained three orders, Asparagales, Liliales and Dioscoreales (which now included the stemonoids). This analysis also allowed for the establishment of a single synapomorphy, although this time by the presence of an inferior ovary. Significantly, the authors noted that it was no wonder the authors of angiosperm classifications had been exasperated by the Lilianae.[27]
Angiosperm Phylogeny Group
These findings, presented at the first Monocot Conference in 1993,[28] with the addition of several studies that had become available in the interim, formed the basis of the 1998 consensus Angiosperm Phylogeny Group (APG) ordinal scheme. Among other things, the Alismatales were expanded and new orders such as Acorales (a placement for Acorus) and Pandanales (which now represented the stemonoids as well as new families) added. While not formally assigning any supraordinal ranks, the classification did recognize an informal grouping of monocot orders as the commelinoids. Otherwise the APG recognized only six monocot orders (Acorales, Alismatales, Asparagales, Dioscoreales, Liliales and Pandanales). The last four were however grouped together in the resulting cladogram and most closely represent the concept of lilioids, although this left some unplaced monocot families, including Corsiaceae and Petrosaviaceae.[29]
Simultaneous with the release of the 1998 APG classification were two events: the publication of Kubitzki's major monograph on the monocots[30] and the Second Monocot Conference. Kubitzki defined superorder Lilianae as all monocots except superorders Commelinae, Alismatanae and the Acoraceae, that is the four orders Asparagales, Liliales, Dioscoreales and Pandanales.[31] The Monocot Conference devoted an entire section to Systematics of the Lilioids[32] and included an update of their previous research by Chase and colleagues.[33] On this occasion the latter felt that there was now enough data to put forward a definitive classification, defining the Lilioids as comprising the four orders placed in Lilianae by Kubitzki. Rudall and colleagues (2002) followed Chase (2000), in using the term "lilioid monocots" and again noting unresolved polytomy between these four orders and the remaining monocot clades (commelinids and Petrosaviaceae), although at that time the Petrosaviaceae were still unplaced.[7][34]
There was now enough new data to justify revising the APG system, and a new classification was issued in 2003. Although this resulted in changes within the orders, it did not affect the relationship between them. Lilioid monocots were discussed but not formally recognized (commelinids, renamed from commelinoids, being the only supraordinal grouping in the monocots to be named) and Petrosaviaceae remained unplaced.
Textbooks and other sources produced in the last century are inevitably based on older classifications. Publications using versions of the APG system are now appearing and the
Phylogeny and evolution
The
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Lilioid monocots 122 |
While this is the most commonly understood relationship, Davis et al. (2013) using a combination of
Subdivision
Five orders make up the lilioid monocots.
- Petrosaviales Takht.(1997)
- Dioscoreales R.Br.(1835)
- Pandanales R.Br. ex J.Presl(1820)
- Liliales Perleb(1826)
- Asparagales Link 1829
PetrosavialesPetrosaviales are a very small order (1 family, 2 genera, about 5 species) of rare leafless achlorophyllous, monosulcate pollen, and follicular fruit.[49]
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Dioscoreales (yams)Dioscoreales are a small order (3 families, 21 genera, about 1,000 species) of mainly contraceptives. They form a sister group to the Pandanales.[50]
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Pandanales (pandans)Pandanales are a medium size order (5 families, 36 genera, about 1,300 species) mainly tropical order many species of which produce strap-like leaves used in the manufacture of baskets, mats and straw hats. The order is very diverse including trees, vines and forest floor saprophytes. They are a sister group to the Dioscoreales.[51] |
Liliales (lilies)Liliales are a large size order (10 families, 67 genera, about 1,500 species) distributed worldwide, particularly in subtropical and pharmaceuticals.[52]
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Asparagales (includes orchids, irises, agaves, amaryllis, and onions)Asparagales are a large very diverse order (14 families, 1,122 genera, about 36,000 species), including many geophytes, ornamental flowers, vegetables, and spices. Most are herbaceous perennials, but some are trees and climbers.[53]
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See also
- Lilianae
- List of systems of plant classification
- List of lilioid families
Notes
- ^ Dahlgren did not actually use the exact term.
- ^ "This superorder is extraordinarily variable and contains some groups which, in our estimation, are likely to have retained many features from the ancestral monocotyledons. The wide range of variation makes a definition difficult and in an evolutionary sense this unit is undoubtedly paraphyletic rather than monophyletic "(emphasis added).[25]
- ^ Subsequent to the 1993 Monocot Conference, and prior to publication.
References
- ^ Zomlefer et al. 2001
- ^ Mondragon-Palomino & Theissen 2009
- ^ Rudall 2002.
- ^ Craene 2010, Lilioids p. 98.
- ^ Botany 400 2016
- ^ Glover 2014, Petaloid monocots p. 122
- ^ a b Furness & Rudall 2001
- ^ a b Johansen et al. 2006
- ^ Lindley 1830, Petaloideae p. 252
- ^ Hutchinson 1936
- ^ a b Heywood 1981
- ^ University of Alberta 1999
- ^ Kron & Chase 1995
- ^ a b Cronquist 1981
- ^ Thorne 1983
- ^ Thorne 1992
- ^ a b Dahlgren, Clifford & Yeo 1985
- ^ Meerow 2002
- ^ Chase & Palmer 1989
- ^ Chase et al. 1993
- ^ Duvall et al. 1993
- ^ a b c APG III 2009
- ^ a b Chase et al. 1995a
- ^ Dahlgren, Clifford & Yeo 1985, p. 94 Fig. 37 Node 8
- ^ Dahlgren, Clifford & Yeo 1985, p. 107
- ^ Dahlgren, Clifford & Yeo 1985, pp. 107–274 Liliiflorae
- ^ Chase et al. 1995b
- ^ Rudall et al. 1995
- ^ APG I 1998
- ^ Kubitzki & Huber 1998
- ^ Kubitzki & Huber 1998, fig. 19 p. 28
- ^ Wilson & Morrison 2000, Systematics of the Lilioids pp. 345–524
- ^ Chase et al. 2000
- ^ Rudall & Bateman 2002
- ^ APG II 2003
- ^ Columbus et al. 2006
- ^ Chase et al. 2006
- ^ Graham et al. 2006
- ^ Chase 2004
- ^ a b Zeng et al. 2014
- ^ a b c d e Hertwick et al. 2015
- ^ Stevens 2016.
- ^ WCLSPF 2015
- ^ a b RBG 2010
- ^ Judd et al. 2007
- ^ a b Chase & Reveal 2009
- ^ a b c Davis et al. 2013
- ^ Hedges & Kumar 2009, p. 205.
- ^ Cameron, Chase & Rudall 2003
- ^ Caddick et al. 2002a.
- ^ Rudall & Bateman 2006.
- ^ Kim et al. 2013.
- ^ Pires et al. 2006.
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- ISBN 978-3-642-64903-5, retrieved 2016-01-21
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- ISBN 978-0-947643-85-0, retrieved 2014-01-14
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- Fay, M. F.; Caddick, L. R.; Cameron, K. M.; Hoot, S., Molecular phylogenetics of Lilianae, vol. 1, pp. 109–137, in Rudall et al. (1995)
- Chase, M. W.; Stevenson, D. W.; Wilkin, P.; Rudall, P. J., Monocot systematics: A combined analysis, vol. 2, pp. 685–730, in Rudall et al. (1995)
- Chase, M.W.; Soltis, D. E.; Soltis, P. S.; Rudall, P. J.; Fay, M. F.; Hahn, W. H.; Sullivan, S.; Joseph, J.; Molvray, M.; Kores, P. J.; Givnish, T. J.; Sytsma, K. J.; Pires, J. C., Higher-level systematics of the monocotyledons: An assessment of current knowledge and a new classification, pp. 3–16, in Wilson & Morrison (2000)
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- Davis, Jerrold I.; Mcneal, Joel R.; Barrett, Craig F.; ISBN 9781139002950, in Wilkin & Mayo (2013)
- Graham, S.W.; Zgurski, J.M.; McPherson, M.A.; Cherniawsky, D.M.; Saarela, J.M.; Horne, E.S.C.; Smith, S.Y.; Wong, W.A.; O'Brien, H.E.; Biron, V.L.; Pires, J.C.; Olmstead, R.G.; Chase, M.W.; Rai, H.S., Robust inference of monocot deep phylogeny using an expanded multigene plastid data set (PDF), pp. 3–21, retrieved 2014-01-04, in Columbus et al. (2006)
- Johansen, Bo; Frederiksen, Signe; Skipper, Martin (2006), "Molecular basis of development in petaloid monocot flowers" (PDF),
- Kron, Kathleen A; ISBN 9780521022897, in Gibbs et al. (1995)
- Articles
- Caddick, Lizabeth R.;
- Cameron, Kenneth M.; S2CID 39771490
- JSTOR 2444471
- JSTOR 2399846
- PMID 21652314
- Duvall, Melvin R.; Clegg, Michael T.; JSTOR 2399849
- Furness, Carol A.;
- Hertweck, Kate L.; Kinney, Michael S.; Stuart, Stephanie A.; Maurin, Olivier; Mathews, Sarah;
- Hutchinson, John (1936), "A new phylogenetic classification of the monocotyledons", Proceedings of the Zesde Internationaal Botanisch Congres, Amsterdam, 2–7 September 1935, ii: 129–131
- Kim, Jung Sung; Hong, Jeong-Ki; Chase, Mark W.; Fay, Michael F.; Kim, Joo-Hwan (May 2013). "Familial relationships of the monocot order Liliales based on a molecular phylogenetic analysis using four plastid loci: matK, rbcL, atpB and atpF-H". Botanical Journal of the Linnean Society. 172 (1): 5–21. .
- Mondragon-Palomino, M.; Theissen, G. (13 January 2009), "Why are orchid flowers so diverse? Reduction of evolutionary constraints by paralogues of class B floral homeotic genes", PMID 19141602
- Pires, J.C.; Maureira, I.J.; Givnish, T.J.; Sytsma, K.J.; Seberg, O.; Petersen, G.; Davis, J. I.; Stevenson, D.W.; Rudall, P.J.; Fay, M.F. & Chase, M.W. (2006), "Phylogeny, genome size, and chromosome evolution of Asparagales", doi:10.5642/aliso.20062201.24)
{{citation}}
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- Thorne, Robert F. (February 1983), "Proposed new realignments in the angiosperms", Nordic Journal of Botany, 3 (1): 85–117,
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- Zeng, Liping; Zhang, Qiang; Sun, Renran; Kong, Hongzhi; Zhang, Ning; Ma, Hong (24 September 2014), "Resolution of deep angiosperm phylogeny using conserved nuclear genes and estimates of early divergence times", PMID 25249442
- Zomlefer, Wendy B.; Williams, Norris H.; Whitten, W. Mark; PMID 21669700.
- APG
- JSTOR 2992015
- Websites
- Department of Botany, University of Wisconsin-Madison, Diversity and Evolution of Monocots: Lilioids (Botany 400 lecture notes), retrieved 2016-01-21
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- Stevens, P.F. (2016) [2001], Angiosperm Phylogeny Website, Missouri Botanical Garden, retrieved 2016-02-01