Taxonomy of Liliaceae

Liliaceae
Lily Family
Temporal range: 68–0Ma

Late Cretaceous - Recent

Lilium candidum
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Monocots
Order: Liliales
Family: Liliaceae
Juss.[1]
Type genus
Lilium
L. Sp. Pl. 1: 302. (1753)[2]
Type species
Lilium candidum
L. Sp. Pl. 1: 302.(1753)[2]
Subfamilies and tribes

sensu APWeb[3]

The taxonomy of Liliaceae has had a complex history since its first description in the mid eighteenth century. Originally the Liliaceae were defined as having a calyx of six equal coloured parts, six stamens, a superior ovary, a single style, and a trilocular capsule. The Liliaceae family progressively expanded until it became the largest of all of the families and extremely diverse being somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary, eventually coming to encompass about 300 genera and 4,500 species. It was thus a 'catch-all' (paraphyletic) taxon. Only since the more modern taxonomic systems, represented by the Angiosperm Phylogeny Group (APG) based on phylogenetic principles, has it been possible to recognize all of its constituents and separate them leaving a relatively small modern family.

The Liliaceae arose in Eurasia, around 68 million years ago with the development of two main evolutionary clades, the Lilileae from the Himalayas, and the Tulipae (Erythronium, Tulipa), (Gagea) from East Asia. Clintonia-Medeola may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae. Divergence amongst the Liliales probably occurred around 36 million years ago. Liliaceae fossils have been dated to the Paleogene and Cretaceous eras in the Antarctic.

Core Liliales probably arose as shade plants, with subsequent evolution of Liliaceae s.s. and Calochortus to open areas including deciduous forest in the more open autumnal period. This was accompanied by a shift from rhizomes to bulbs, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves.

While the suprageneric structure of the family has varied greatly with its ever changing circumscription, as currently constituted it consists of three subfamilies, of which the Lilioidea is further divided into tribes, with fifteen genera and approximately 600 species of flowering plants.

History

Pre-Darwinian

The type genus, Lilium, from which the name of the family was derived, was described by Carl Linnaeus in 1753, with seven species.[2] He placed Lilium within the Hexandria Monogynia (six stamens, one carpel) in his sexual classification in the Species Plantarum.[4]

The Liliaceae family was first described by Michel Adanson in 1763,[5][6] but formally named by Antoine Laurent de Jussieu in 1789.[1] Adanson described eight subfamilies with seventy eight genera, however the subfamily he described as Lis (Lilies) had seven genera (Uvularia, Mithridatium, Mendoni, Lilium, Imperialis (Fritillaria), Fritillaria and Tulipa) of which four are in the modern genus.[7] De Jussieu placed them as the fourth Ordo (Order: Lilia) of the third class (Stamina Perigyna) of Monocots, with eight genera (Tulipa, Erythronioum, Methonica, Uvularia, Fritillaria, Imperialis, Lilium, Yucca) only four of which remain in the family. However the use of the term Ordo at that time was closer to what we now understand as Family, rather than Order.[8][9] Although Jussieu used the term 'Lilia' in his Genera Plantarum, elsewhere he used the French term 'Liliacées',[10] as had Adanson before him using both French and Latin, and the word 'Liliaceae' was soon widely used by botanists such as Samuel Frederick Gray,[11] John Lindley,[12] and Pierre-Joseph Redouté[13] in the early nineteenth century. Jussieu defined this grouping as having a calyx of six equal coloured parts, six stamens, a superior ovary, single style, and trilocular capsule. Gray (1821) provided the first description of Jussieu's scheme in English, identifying two genera occurring in Britain (Tulipa, Fritillaria), distinguished by the absence or presence of basal nectaries. His key used the presence of six stamens, a single style, equal stamina, a simple petaloid perianth and a trilocular capsule with flat seeds to identify the family.[14] Although De Candolle (1813) had not explicitly described the Liliaceae, his overall classification scheme influenced many later writers including Gray.[15] In this scheme,[16] the Liliaceae were considered a family within the Endogenæ phanerogamæ, that is those vascular plants (Vasculares) whose vascular bundles were thought to arise from within (Endogènes, endogenous), a term he preferred to monocots (Monocotylédonés). Of these, the true monocots of Jussieu became the Phanerogams or Phanérogames (Phenogamae in Gray), hence Endogenæ phanerogamæ.[17] Candolle also instituted the concept of ordered ranks, based on classes, subclasses, familles (Latin: ordines naturales) and tribus (tribes),[9] subdividing the Liliaceae.

Lindley was the first English systematist, publishing his work in 1830,[18] and following the reasoning of Jussieu he used the term tribe to describe the Liliaceae as a division of the hexapetaloid monocots, characterised by a superior ovary, highly developed perianth, inward turning anthers, a trilocular polyspermous capsule and seeds with a soft spongy coat. He offered seven genera as examples (Erythronium, Lilium, Calochortus, Blandfordia, Polianthes, Hemerocallis and Funkia). By 1846, in his final work, he refined and greatly expanded his taxonomy further favouring the term Alliances of Endogens over monocots, as a class, with eleven alliances including the Liliales. This includes four Orders (families) including Liliaceae which he referred to as lilyworts in the vernacular. In this work he unhappily acknowledged the great diversity in the circumscription of the order, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already paraphyletic ("catch-all"), being all Liliales not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. He recognised 133 genera and 1200 species.[19] By the time of the next major British classification, that of Bentham and Hooker in 1883 (published in Latin) several of Lindley's other families had been absorbed into the Liliaceae.[20]

Post-Darwinian

Although the appearance of Charles Darwin's Origin of Species in 1859 changed the way that taxonomists considered plant classification, incorporating evolutionary information into their schemata, this did little to further define the circumscription of Liliaceae.[21] For instance, the first phylogenetic based system, by Bessey (1915) followed Engler in defining Liliaceae as "Pistil mostly 3-celled; stamens 6; perianth of two similar whorls, each of three similar leaves", although placing the Liliales in a novel sublass of monocots, the Strobiloideae.[22] The Liliaceae were treated as Liliiflorae by Wettstein (1901–1935) and by Engler and Prantl (1931), and Liliales from the systems of Hutchinson (1959) onwards. Over time the Liliaceae became increasingly broadly, and somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary, eventually coming to encompass about 300 genera and 4,500 species, within the order Liliales in the scheme of Arthur Cronquist (1981) who merged it with the Amaryllidaceae.[23] Later similar schemes include the system of Robert F. Thorne[24] and that of Armen Takhtajan.[25] They were characterised as petaloid monocots, characterized by showy flowers with tepals and without starch in the endosperm. Cronquist had placed most flowering petaloid monocots with six stamens in this very broad (and clearly polyphyletic) family.[26] Cronquist rejected the importance of the position of the ovary and thus included the Amarylloidaceae some species of which had an inferior ovary, and which others separated into a distinct family,[27] although this then created other problems.[28] The Liliaceae were one of the major families in the Cronquist system which considerably broadened the circumscription to include 22 families in addition to Liliaceae s.s..[29]

Dismembering Liliaceae

Other botanists in the twentieth century echoed Lindley's concerns about the phylogeny of the Liliaceae. The earliest of these was Johannes Paulus Lotsy in 1911,[30] followed by Huber in 1969 who suggested four separate families.[31] These various proposals to separate small groups of genera into more homogeneous families, made little impact till that of Dahlgren (1985) incorporating new information including synapomorphy.[26] Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly Liliales and Asparagales).[32][33] Over the 1980s, in the context of a more general review of the classification of angiosperms, the Liliaceae were subjected to more intense scrutiny. By the end of that decade, the Royal Botanic Gardens at Kew, the British Museum of Natural History and the Edinburgh Botanical Gardens formed a committee to examine the possibility of separating the family at least for the organization of their herbaria. That committee finally recommended that 24 new families be created in the place of the original broad Liliaceae, largely by elevating subfamilies to the rank of separate families.[34]

The 1990s saw considerable progress in plant phylogeny and phylogenetic theory, enabling a phylogenetic tree to be constructed for all of the flowering plants. The establishment of major new clades necessitated a departure from the older but widely used classifications such as Cronquist and Thorne based largely on morphology rather than genetic data. This complicated discussion about plant evolution and necessitated a major restructuring.[35] rbcL gene sequencing and cladistic analysis of monocots had redefined the Liliales in 1995[36] from four morphological orders sensu Dahlgren. The largest clade representing the Liliaceae, all previously included in Liliales, but including both the Calochortaceae and Liliaceae sensu Tamura. This redefined family, that became referred to as core Liliales, but corresponded to the emerging circumscription of the Angiosperm Phylogeny Group (1998).[37]

Modern APG classification

To meet this need, flowering plant systematists formed the Angiosperm Phylogeny Group (APG) resulting in a new classification published in 1998[38] largely based on the work of Kåre Bremer and colleagues at Uppsala and Stockholm universities in the late 1970s, [39][40][41] which became available on the internet in 1996.[42] This scheme was ordinal (concentrating on orders rather than families), prioritising monophyly. However progress was rapid and the modern era of the taxonomy of the Liliaceae family comes from Judd et al.[28] (second edition 2002), the APG II (2003[35]) and APG III (2009 [43]), while the Linear APG III assigned it the family number 61. [44] While the original APG did not specifically address the issues of the polyphyly within Liliaceae, APG II did so within the two closely morphologically related orders, Liliales and Asparagales[45] recognising the continued common use of the sensu lato (s.l.).

These studies of DNA and morphological data (particularly reproductive morphology) together with phylogenetic analyses, allowed the conclusion that the "petaloid monocots" do not really belong to one botanical family but rather are distributed across two separate different orders, the Asparagales and Liliales. The monophyly of these newly defined orders is supported by cladistic analysis based on morphology, 18S rDNA, and many other DNA sequences, particularly the plastid gene rbcL.[46][47][48][49][50][51][52][53][54][55][56][57][58][59][60][61][62][63][64]

Together with other analyses within each of these two orders has allowed the redistribution of the original genera of Liliaceae sensu lato into a variety of families, as illustrated in the two cladograms of the Asparagales and Liliales shown here.[65] These changes have resulted in considerable changes in the positioning of Liliaceae within the overall classification, as shown in the table below, as well as within the family.

Table 1: Evolution of placement of Liliaceae in different taxonomic schemes[21]
Rank Bentham and Hooker (1883)[20]Cronquist (1981)[23]Takhtajan (1980)[66] Dahlgren (1985)[26]Thorne (1992)[24] APG (2003-9)[35][43]
Division MagnoliophytaMagnoliophyta
ClassMonocotyledonsLiliopsidaLiliopsidaLiliopsidaAngiospermae
Subclass LiliidaeLiliidaeLiliidaeLiliidae
Superorder (Series) Coronarieæ DioscoreanaeLiliifloraeLilianaeMonocots
Order LilialesLilialesLilialesLilialesLiliales

For a comparison of the classifications of Genera from 1959 (Hutchinson)[67] to 2000 (Wilson and Morrison)[68] see Table 1 in Fay et al. 2006.[69] and Table 1 in Peruzzi et al. 2009[70]

Cladistic analysis of Liliales and families within former Liliaceae

Within the Liliales, cladistic analysis revealed the Liliaceae as one of four main groupings.[45] The following cladograms based on molecular phylogenetics reveal the polyphyletic nature of the original Liliaceae family, which encompassed a large number of differing groups of genera. This led to botanists increasingly adopting a stricter more narrow monophyletic concept (sensu stricto) of the family based on phylogenetic relationships, as expressed in the 2009 APG III system, rather than the older broader one (sensu lato),

Asparagales

Orchidaceae

Hypoxidaceae (traditionally placed in Amaryllidaceae)

Iridaceae
Xanthorrhoeaceae sensu lato

Hemerocallidaceae (traditionally placed in Liliaceae)

Xanthorrhoeaceae (traditionally placed in Liliaceae)

Asphodelaceae (traditionally placed in Liliaceae)
Alliaceae sensu lato

Agapanthaceae (traditionally placed in Liliaceae)

Alliaceae (traditionally placed in Liliaceae)

Amaryllidaceae


Laxmanniaceae (traditionally placed in Liliaceae)

Asparagaceae (traditionally placed in Liliaceae)

Ruscaceae (traditionally placed in Liliaceae)


Themidaceae (traditionally placed in Liliaceae)

Hyacinthaceae (traditionally placed in Liliaceae)

Agavaceae (traditionally placed in Amaryllidaceae or Liliaceae)








Liliales


Alstroemeriaceae (traditionally placed in Liliaceae)

Colchicaceae (traditionally placed in Liliaceae)

Melanthiaceae (traditionally placed in Liliaceae)

Smilacaceae (traditionally placed in Liliaceae)

Liliaceae sensu stricto




Other displaced groups included members of the Nartheciaceae, Tecophilaeaceae, Tofieldiaceae, and Uvulariaceae families and Dioscoreales, and Alismatales orders.

Changes to family structure in APG III

The APG III system when it was published in 2009, greatly expanded the families Xanthorrhoeaceae, Amaryllidaceae, and Asparagaceae.[71] Thirteen of the families of the earlier APG II system were thereby reduced to subfamilies. The APG II families and their equivalent APG III subfamilies are as follows:

Phylogeny

Sequencing of the rbcL and matK regions of the chloroplast DNA of Lilium and related genera[72] confirmed the circumscription of the family s.s. as described by Tamura (1998).[73] Chloroplast ndhF gene sequencing also supports Liliaceae monophyly, reuniting the Liliaceae and Calochortaceae sensu Tamura.[37][69]

Despite establishing this relative degree of monophyly for the Liliaceae family,[36][74] morphology remains diverse[73] and there exists within the Liliaceae clade, a number of subclades. Particularly enigmatic were Clintonia, Medeola, Scoliopus, and Tricyrtis. Clintonia (which has a disjunct distribution involving both East Asia and North America) and the closely related Medeola form one of those subclades, and are now considered a separate tribe (Medeoleae), within the Lilioideae although at different times they have been both a separate subfamily (Medeoloideae) and family (Medeolaceae). Sequencing of the rbcL and matK chloroplast genes established monophyly for Clintonia, but with separate clades corresponding to the two distribution regions.[75]

While the ten genera of the Lilioideae subfamily are characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores), the five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group, previously characterised under the Calochortoideae alone. They are characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac. At times these genera were considered in a separate family (Calochortaceae) (e.g. Tamura) or even the more heterogeneous Uvulariaceae sensu Dahlgren. However most of the latter had low morphological similarity to the Liliaceae and Uvularia and Disporum are now located in the Colchicaceae. Disporum contained both Asian and North American species which had always been distinguishable. Following molecular analysis the North American species were restored to the genus Prosartes and retained in Liliaceae, and are now included in the Streptopoideae. The Asian species were moved to Colchicaceae.[76]

Evolution

The development of a phylogenetic approach to taxonomy, starting with Charles Bessey's The phylogenetic taxonomy of flowering plants (1915) suggested the Liliales formed some of the earliest monocots.[22] Molecular analysis suggests that Liliaceae arose in Eurasia, around 68 million years ago during the late (Maastrichtian) Cretaceous to early (Paleocene) Paleogene eras[77] with the development of two main evolutionary clades. The first of these, characterised as Lilileae (Lilium, Fritillaris, Nomocharis), Cardiocrinum), Notholirion) from the Himalayas, and the second, the Tulipae (Erythronium, Tulipa), (Gagea) from East Asia. On the other hand Clintonia-Medeola may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae sensu Tamura. Divergence amongst the Liliales probably occurred around 36 million years ago, with the Liliaceae (sensu Tamura) emerging at 27 million years, Liliaceae s.s. at 20 million, Lilieae 12 million and Calochortus 7 million.[37] Liliaceae fossils have been dated to the Paleogene[78] and Cretaceous[79] eras in the Antarctic.

Core Liliales probably arose as shade plants, with subsequent evolution of Liliaceae s.s. and Calochortus to open areas including deciduous forest in the more open autumnal period, but then a return of some species (e.g. Cardiocrinum). This was accompanied by a shift from rhizomes to bulbs, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves. Again, some reversal to the broader reticulate-veined leaves occurred (e.g. Cardiocrinum) [37]

Subdivisions and genera

Leaves of fruit of Clintonia borealis (Medeoleae) growing in Canada
Flower of Nomocharis aperta (Lilieae) growing in China
Flower of Calochortus catalinae (Calochortoideae) growing in California
Leaves and berries of Streptopus lanceolatus, North America (Streptopoideae)

Suprageneric subdivisions

Due to the diversity of the originally broadly defined Liliaceae s.l., many attempts have been made to form suprageneric classifications, e.g. subfamilies and tribes.[70] By 1813, Candolle recognised five subdivisions which he called tribes (Asparagées, Trilliacées, Asphodelées, Bromeliées, Tulipacées),[16] all of which Jussieu had made separate families, with the exception of Tulipa, which was a genus within the Liliaceae. By 1845 John Lindley observed that the family had become extremely diverse, ill defined and unstable not only by its overall circumscription, but also by its subdivisions. Of the 133 genera he included, he described eleven suborders.[19]

In 1879 a revision of the North American Liliaceae described sixteen tribes[80] and Bentham and Hooker twenty tribes in 1883, while Engler and Prantl in their extensive description of the Liliaceae in 1899 identified 31 tribes distributed over 11 subfamilies.[81] In 1936 Buxbaum undertook a major revision of the Liliaceae and inter alia described the subfamily Lilioideae, with three tribes: Lloydieae, Tulipeae and Lilieae.[82][83][84] Peruzzi et al. (2009) discuss the complex rearrangement of the various genera, tribes and subfamilies over a 30 year period (Peruzzzi et al. Table1) from 1985,[70] partly summarised here in Table 2.

Classifications published since the use of molecular methods in phylogenetics have taken a narrower view of the Liliaceae. In 1998 Tamura considered Calochortus sufficiently distinct to elevate the subfamily Calochortoideae to family status as Calochortaceae.[73][85] In 2009, Takhtajan used a very narrow definition (see the table below).[66] As of January 2014 the Angiosperm Phylogeny Website (APweb)[3] included four of Takhtajan's families in its Liliaceae, recognizing three subfamilies, one of which is divided into two tribes.[86]

Table 2: Comparison of Four 21st Century Classifications of the Liliaceae
Tamura[73] Takhtajan[66] Taxonomicon[87] APweb[86]
Family Subfamily Tribe Family Tribe Family Subfamily Tribe Family Subfamily Tribe
Liliaceae Lilioideae Lilieae Liliaceae Lloydieae Liliaceae Lilioideae Lloydieae Liliaceae Lilioideae Lilieae
Lilieae Lilieae
Tulipeae Tulipeae Tulipeae
Medeoloideae Medeolaceae Medeola, Clintonia Medeoleae
Calochortaceae Calochorteae Scoliopaceae Calochortoideae Calochortoideae
Tricyrtideae Tricyrtidaceae Streptopoideae Streptopoideae

Genera

History

The Cronquist system (1981) had nearly 300 genera in Liliaceae. Most of those have been reassigned to other families, as shown here, following ITIS, together with their disposition as APG III transfers to other families (including subfamilies and orders other than Liliales where applicable) in brackets (Family: Subfamily, Order). Current members of Liliaceae in bold:

(Asian species. N American species restored as Prosartes D. Don)
(Asparagaceae: Nolinoideae, Asparagales)

The following includes other genera that have historically been classified in family Liliaceae s.l.:

Modern classification

Various authorities (e.g. ITIS 16,[88] GRIN 27,[89] WCSP,[90] NCBI,[91] DELTA[92] ) differ on the exact number of genera included in Liliaceae s.s., but generally there are about fifteen to sixteen genera, depending on whether or not Amana is included in Tulipa and Lloydia in Gagea. For instance Amana is still listed separately in WCSP.

Currently the APWeb lists fifteen genera, arranged as follows:[3][93]

The largest genera are Gagea (200), Fritillaria (130), Lilium (110), and Tulipa (75 species), all within the Lilieae tribe. For a description of the phylogenetic relationships between each of the genera, see Fig. 2.7 in Kamenetsky and Okubo (2012).[94]

Etymology

From Lilium the type species which is Latin for Lily, which in turn came from the Greek λείριον (leírion).

References

  1. ↑ 1.0 1.1 Jussieu 1789, Lilia pp. 48–49.
  2. ↑ 2.0 2.1 2.2 Linnaeus 1753, Lilium vol. i p. 302.
  3. ↑ 3.0 3.1 3.2 Stevens 2015, Liliaceae.
  4. ↑ Linnaeus 1753, Hexandria monogynia vol. i pp.  285–352.
  5. ↑ Lobstein 2013.
  6. ↑ Adanson 1763, VIII Liliaceae p. 42–60.
  7. ↑ Adanson 1763, Liliaceae Section II Les Lis Lilia p. 48.
  8. ↑ ICN 2011, 18.2 Names of families and subfamilies, tribes and subtribes.
  9. ↑ 9.0 9.1 A. P. de Candolle 1813, Des familles et des tribus pp. 192–195.
  10. ↑ de Jussieu 1778, p. 228.
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  14. ↑ An Arrangement of the Families, and of the anomalous Genera of phenogamous Plants, according to their sexual Organs. p. vol 2: vi. Retrieved 4 February 2014. In Gray (1821).
  15. ↑ The genera of British plants, according to their mutual relations, with the number of species in each genus. 2. Plantae Endogenae: B Plantae Endogenae Phenogamae - 14. Liliaceae. p. vol 1: xx. Retrieved 5 February 2014. In Gray (1821).
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  34. ↑ Mathew, Brian (1989). "Splitting the Liliaceae". The Plantsman 11 (2): 89–105.
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  37. ↑ 37.0 37.1 37.2 37.3 Patterson, T. B.; T. J. Givnish (2002). "Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from rbcL and ndhF sequence data" (PDF). Evolution 56 (2): 233–252. doi:10.1111/j.0014-3820.2002.tb01334.x. PMID 11926492. Retrieved 14 January 2014.
  38. ↑ APG (1998). "An ordinal classification for the families of flowering plants". Annals of the Missouri Botanical Garden. 85: 531-553. (4): 531–553. JSTOR 2992015.
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  40. ↑ Bremer, K.; Bremer, B.; Thulin, M. (1995) [1996, 1997]. Introduction to Phylogeny and Systematics of Flowering Plants. 1st, 2nd, 3rd eds. Uppsala University.
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Bibliography

Systematics

Taxonomic classifications

Table of 58 families, Part II: Page 1
Table of 1615 genera, Part II: Page 8

Symposia

Databases

External identifiers for Liliaceae
Encyclopedia of Life 4174
GBIF 7699
ITIS 42633
NCBI 4677
Also found in: Wikispecies, Arctos,Paleobiology Database
  • The dictionary definition of Liliaceae at Wiktionary
  • Media related to Liliaceae at Wikimedia Commons