LIGNOPHYTA

True roots +; lateral meristems: cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially.

EXTANT SEED PLANTS/SPERMATOPHYTA

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins derived from (some) sinapyl and particularly coniferyl alcohols, thus containing p-hydroxyphenyl and guaiacyl lignin units, (lignins derived from p-coumaryl alcohol, i.e. S [syringyl] lignin units); true roots present, apex multicellular, xylem exarch, and branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, plastids with starch grains; phloem fibres +; stem cork cambium superficial, root cork cambium deep seated; leaves with single trace from sympodium ["nodes 1:1"]; stomata ?; leaf vascular bundles collateral; leaves megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, lamina with vein density up to 5 mm/mm² [mean for all non-angiosperms 1.8]; axillary buds associated with at most some leaves; prophylls [including bracteoles] two, lateral; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, developing after pollination, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], S [syringyl] lignin units common, positive Maüle reaction [syringyl:guaiacyl ratio more than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma +; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cells from same mother cell that gave rise to the sieve tube; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves petiolate, lamina [formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, polysymmetric, parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P not sharply differentiated, with a single trace, outer members not enclosing the rest of the bud, often smaller than inner members; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], ± embedded in the filament, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not secretory]; ovules few [?1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, megaspore tetrad linear, functional megaspore chalazal, lacking sporopollenin and cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; P deciduous in fruit; seed exotestal; pollen binucleate at dispersal, trinucleate eventually, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing at 80-600 µm/hour, with pectic outer wall, callose inner wall and callose plugs, growing between cells, penetration of ovules via micropyle [porogamous] within ca 18 hours, distance to first ovule 1.1.-2.1 mm, tube moves between nucellar cells; double fertilisation +, endosperm diploid, cellular [micropylar and chalazal domains develop diffently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, ndhB gene 21 codons enlarged at the 5' end, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and three copies of the PHY gene, [PHYB [PHYA + PHYC]].

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable homoplasy as well as variation within and between families of the ANITA grade in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous... For other features such as details of sugar transport in the phloem, their placement on the tree is frankly speculative. Finally, for features such as parietal tissue/a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), I am unsure where on the tree a thicker nucellus and a stylar epidermal layer are acquired.

[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: vessels +, elements with elongated scalariform perforation plates; wood fibres +; axial parenchyma diffuse or diffuse-in-aggregates; tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

LILIALES Perleb  Main Tree, Synapomorphies.

Plants geophytes; stem fructans, chelidonic acid, steroidal saponins +; (velamen +); (cuticular waxes as parallel platelets); leaves elliptical, (main veins seven or fewer), fine venation reticulate, leaf base not sheathing; inflorescence terminal; T large, (spotted), with tepal nectaries; anthers extrorse; style long, stigma capitate; ovules many/carpel, parietal tissue absent, nucellar cap +; P deciduous; tegmen with cellular structure; endosperm with thick pitted walls, hemicellulosic; mitochondrial sdh3 gene lost. - 11 families, 67 genera, 1558 species.

Evolution. Divergence & Distribution. Stem group Liliales are dated to ca 124 million years before present, crown group Liliales to ca 117 million years before present (Janssen & Bremer 2004), rather older than the estimates in Bremer (2000); comparable figures in Magallón and Castillo (2009) are ca 135.7 (stem) and 125.5 (crown) and 120.1 (stem) and 114.4 (crown) million years for relaxed and constrained penalized likelihood datings respectively.

Genes & Genomes. Very large genomes with a C value of some 35 picograms or more are found in some Melanthiaceae, Liliaceae and Alstroemeriaceae (Leitch et al. 2005), while Paris japonica (Melanthiaceae), at over 150 picograms, has the largest known genome of any organism (Pellicer et al. 2010a).

Chemistry, Morphology, etc. Asparagales commonly have Arum-type arbuscular mycorrhizae, while in Liliales such mycorrhizae are commonly Paris-type (see F. A. Smith & Smith 1997). Three-trace tepals occur in some taxa here including Corsiaceae (Rudall and Eastman in Neyland 2002), Melanthiaceae, Liliaceae, and Colchicaceae, but not in some Asparagales and also Petermanniaceae - at what level is this a phylogenetically interesting character? See Endress (1995b) for dome details of floral development. Glucomannan seed reserves are reported from some species of Liliaceae-Lilioideae and Colchicaceae-Colchiceae (Jakimow-Barras 1973). For chromosome size of a number of taxa in the group, see Vijayavalli and Mathew (1990 - as Liliaceae) and Tamura (1995).

There is much information in Rudall et al. (2000), including a summary of pollen variation; for pollen of Japanese representatives, see Handa et al. (2001). Some details of ovule morphology, etc., are taken from Oganezova (2000b); see El-Hamidi (1952) for gynoecium and Conran et al. (2009b) for information on leaf morphology and stomata.

Phylogeny. For phylogenetic relationships, see Chase et al. (1995a), Rudall et al. (2000), Patterson and Givnish (2002), Vinnersten and Bremer (2001), Chen et al. (2007: Bayesian analysis, support for many branches weak) and especially Fay et al. (2006c). The tree here is based on this last study, but there are substantial areas of uncertainty which may yet necessitate recognising formally more clades. Neyland (2002a), analysing variation in 26S ribosomal DNA, suggests that Corsiaceae are to be associated with Liliales. Although this position has only weak support, it is largely consistent with morphological evidence; Davis et al. (2004) also found Corsiaceae to be associated here. Davis et al. (2004) did not find Smilaceae and Rhipogonaceae to be sister taxa, although they are so similar (combining the two is even an option in A.P.G. [2003]), but there was no strong support for this (non!) position, and sampling was poor. However, Fay et al. (2006c: weak support), Givnish et al. (2006: strong support), and Chase et al. (2006) all find Philesiaceae and Rhipogonaceae to be sister taxa; Smilacaceae are strongly supported as sister to Liliaceae. Relationships suggested by the rbcL study of Janssen and Bremer (2004) are rather different, but neither Petermanniaceae nor Corsiaceae were included.

Previous Relationships. Cronquist (1981) circumscribed Liliales very broadly, and his order now makes up the bulk of Liliales and Asparagales.

Includes Alstroemeriaceae, Campynemataceae, Colchicaceae, Corsiaceae, Liliaceae, Melanthiaceae, Petermanniaceae, Philesiaceae, Rhipogonaceae, Smilacaceae.

Synonymy: Alstroemeriales Hutchinson, Campynematales Doweld, Colchicales Dumortier, Liriales K. Koch, Melanthiales Link, Paridales Link, Smilacales Link, Trilliales Takhtajan, Veratrales Dumortier

CORSIACEAE Beccari, nom. cons.   Back to Liliales

Echlorophyllous mycoheterotrophic herbs; chemistry?; vessels?; root endodermis not obvious; epicuticular wax platelets parallel, stomata ?type; leaves spiral to two-ranked, sheath closed, ?main veins; (plant dioecious); flowers single, monosymmetric, T large, median T of outer whorl adaxial and standard-like [= "labellum"], ?nectary [callus on "labellum" in Corsia?], (A basally adnate to style [gynostemium] - Corsia; A 5 + staminode opposite "labellum"), pollen porate, ovary inferior, placentation also parietal, style short, (unbranched - Corsia), or stigma sessile; ovules many/carpel, integuments two cells across, parietal tissue 1 cell across, nucellar cap?, funicle long; fruit capsular [type?], or with terminal pore; seeds minute, testal; (endosperm also with starch), embryo undifferentiated; n = 9; seedling?

3[list]/30. S. China, S. South America, Papuasia-Australia (map: from van Royen 1972; Ibisch et al. 1996). [Photo - Flower]

• Corsiaceae are echlorophyllous mycoheterotrophic herbs that have leaves with closed sheaths and large, single, terminal monosymmetric flowers that are held in the morphologically "inverted" (for monocots) position; the median member of the outer whorl, the labellum (sic), is much larger than all other tepals and is adaxial. The six stamens are extrose and the ovary is inferior.

Evolution. Floral Biology. The flowers of most species of Corsia are inverted and the "labellum" is held horizontally, the stamens, etc., being underneath it, however, the flowers of C. dispar are held at 180^o to them so the relationships of the parts is inverted (Jones & Gray 2008).

Chemistry, Morphology, etc. For information, see Rübsamen (1986: general the megaspore mother cell seems to have two cell layers above it - ?a nucellar cap), Neinhuis and Ibisch (1998: general), and Rudall and Eastman (2002: morphology).

Previous Relationships. Analysis of 26S rDNA sequences suggested that Corsiaceae were polyphyletic; Arachnitis may be sister to Thismia and/or Burmannia (Burmanniaceae-Dioscoreales - see Neyland & Hennigan 2003: G. Petersen et al. 2006b, combined analysis). These relationships have not been confirmed.

CAMPYNEMATACEAE Dumortier   Back to Liliales

Fibrous leaf bases persistent; chemistry?; stomata type?; leaf lacking reticulate fine venation, base sheathing [?type]; flowers medium-sized, T green, not spotted; A adnate to base of T, tapetum 2- or multinucleate; G ± inferior, styluli +; ovules 3-many/carpel, crassinucellate; fruit a capsule or indehiscent, T enlarging, persistent; seeds angled, exotestal, with phlobaphene; endosperm development?; embryo minute; n = 11, chromosomes to 3µm long; seedling?

2[list]/4. New Caledonia and Tasmania (map: from van Balgooy 1984).

• Campynemataceae are herbs which have medium-sized flowers with green, unspotted tepals that are accrescent in fruit; the ovary is inferior and the styluli are separate.

Evolution. Divergence & Distribution. Stem group Campynemataceae are dated to ca 117 million years before present, crown group Campynemataceae to ca 73 million years before present (Janssen & Bremer 2004: note that Campynemataceae are sister to the rest of Liliales there).

Chemistry, Morphology, etc. Campynemanthe has a subumbellate inflorescence, introrse anthers (see illustration in Kubitzki 1998b) partly superior ovary, and dentate leaf apex; Campynema has extrorse anthers, multinuclear tapetal cells and an inferior ovary.

Additional information is taken from Dahlgren and Lu (1985: Campynemanthe), Lowry et al. (1987: cytology and embryology), and Kubitzki (1998b: general).

[[Petermanniaceae [Colchicaceae + Alstroemeriaceae]], Melanthiaceae, [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]: (fruit a berry).

[Petermanniaceae [Colchicaceae + Alstroemeriaceae]]: primary root of seedling well developed.

Petermannia is apparently sister to the other two families above (J. Davis in Vinnersten & Reeves 2003; Graham et al. 2005), the original sample that placed it inside Colchicaceae (Rudall et al. 2000) being misidentified (see Chase et al. 2006).

PETERMANNIACEAE Hutchinson, nom. cons.   Back to Liliales

Plant climbering, rhizome woody; saponins 0; velamen +; vessels also in stem and leaf; stem with prickles and leaf-opposed tendrils; leaves spiral, subpetiolate, with midrib and 5-7 main veins, secondary veins parallel; inflorescence cymose, pedicels not articulated; T 1-veined, medium-sized; tapetum amoeboid; ovary inferior, placentation parietal, stigma wet; ovules many/carpel, micropyle?; fruit a berry; seed with thickened exo- and endotesta, several-layered mesotesta, a cuticle, and crushed tegmen; n = 5; first leaves cataphylls.

1/1: Petermannia cirrosa. Central part of the E. coast of Australia, rare.

• Petermanniaceae are rhizomatous woody climbers with petiolate leaves that have reticulate venation and leaf-opposed tendrils. The smallish flowers, borne in cymose inflorescences, have an inferior ovary and the fruits are fleshy.

Evolution. Divergence & Distribution. Fossil Petermanniopsis is reported from the early Eocene of Australia (Conran & Christophel 1999).

Chemistry, Morphology, etc. Petermanniopsis has paracytic stomata - probably plesiomorphic (Conran & Christophel 1999). The inflorescences and tendrils are terminal, but become leaf-opposed as they are evicted by the strong growth of an axillary shoot.

Information is taken from Tomlinson and Ayensu (1969: anatomy), Björnstad (1970: embryology), Conran and Clifford (1998: general) and Prychid and Rudall (1999).

Previous relationships Although in early versions of this site and in A.P.G. (2003) Petermanniaceae were included in Colchicaceae, not only are they phenetically easily distinguishable, in particular Petermanniaceae lack the distinctive U-shaped tepals of Colchicaceae and are vegetatively very different. Molecular data now separate them.

[Colchicaceae + Alstroemeriaceae]: ?

Chemistry, Morphology, etc. For fructose oligosaccharide accumulation (only one record for Alstroemeriaceae s.l.), see Pollard (1982).

Phylogeny. Colchicaceae, Alstroemeriaceae (and Luzuriagaceae, here included in the latter family) are commonly linked, e.g. Tamura et al. (2004a), Davis et al. (2004), Janssen and Bremer (2004), Fay et al. (2006c).

COLCHICACEAE de Candolle, nom. cons.   Back to Liliales

Flavones, protoalkaloid colchicine +, steroidal saponins 0; raphides 0; cuticular wax with parallel platelets; leaves two-ranked to ± verticillate, conduplicate, midrib common; inflorescence various; T towards base U-shaped and folded around each stamen in bud, connate or not, (A latrorse, introrse), sexine thick, (pollen operculate); G ([2, 4]), style branched or not, stigma with recurved lobes, wet or dry; ovules 2-many/carpel, ± ascending, orientation various, (unitegmic), micropyle bistomal; antipodal cells multinucleate, raphides 0; seeds rounded (subangled), strophiole or aril +; exotesta with thick walls (less so - Colchicum), phlobaphene +; endosperm nuclear, (with starch), embryo small; chromosomes 1-16 µm long; cotyledon photosynthetic or not, bifacial (ligulate).

15[list]/245 - six tribes below. Temperate to tropical, but not in South America (map: see Meusel et al. 1965; Hong 1993; Nordal et al. 2001; del Hoyo & Pedrola-Monfort 2006, 2008).

1. Burchardieae J. C. Manning & Vinnersten

Rhizome short, vertical, with papery scales; leaves sheathing; styluli +; capsule septicidal.

1/?. Australia.

Synonymy: Burchardiaceae Takhtajan

[Uvularieae [Tripladenieae [Colchiceae [Iphigenieae + Anguillarieae]]]]: flowers axillary.

2. Uvularieae Meisner

Rhizomes +; leaves distichous, (pseudopetiole +); pollen disulcate; nucellar cap 0; (fruit a berry; loculicidal capsule); n = 6-9, chromosomes 5-16 µm long.

2/15: Disporum (10). W. and E. North America, East Asia to W. Malesia.

Synonymy: Uvulariaceae Kunth, nom. cons.

[Tripladenieae [Colchiceae [Iphigenieae + Anguillarieae]]]:

3. Tripladenieae Vinnersten & J. C. Manning

Rhizome +; leaves distichous, (pseudopetiole +); n = 7, 18.

3/5. Australia and New Guinea.

[Colchiceae [Iphigenieae + Anguillarieae]]: tunicated corm +; alkaloids with a 7-C tropolone ring +; (vessels in stem); leaf base sheathing; nectary on filament bases; (antipodal cells persistent).

4. Colchiceae Reichenbach

(Vessels in stem - Sandersonia), (leaves whorled); (tepals connate); nectaries median on tepal or on stamen base (0); (pollen polyporate); antipodal cells divide; n = 6-10, etc., chromosomes 2.8-14.3 µm long

5/170: Colchicum (100, if inc. Androcymbium, 150), Gloriosa (inc. Littonia: 9). Africa, Europe, Central to tropical South East Asia.

Synonymy: Bulbocodiaceae R. A. Salisbury, Merenderaceae Mirbel

[Iphigenieae + Anguillarieae]: ?

5. Iphigenieae Hutchinson

Nectaries 0; n = 10, 11, 13.

2/10: Iphigenia (9). Old World Tropics, South Africa.

6. Anguillarieae D. Don

Inflorescence racemose or spicate, bracts 0; (tepals connate); n = 5, 7, 10, 11.

2/38: Wurmbea (37). Africa, Australia.

• Colchicaceae are cormose or rhizomatous geophytes that may be recognised by their rather large flowers with six, usually free tepals, each more or less enveloping a stamen and with nectaries on the adaxial surface at or towards the base, and a superior ovary. [Photos - Collection].

Evolution. Divergence & Distribution. Stem group Colchicaceae are dated to ca 76 million years before present, crown group Colchicaceae to ca 44 million years before present (Janssen & Bremer 2004). The Colchicum-Androcymbium clade may have diverged from the rest 13.4±1.5 million years ago, probably in southwestrn Africa (del Hoyo et al. 2009).

Ecology & Physiology. Colchicaceae are well represented in the taxa that have water-catching leaves with very distinctive morphologies that are particularly prominent in the foggy deserts of Namaqualand (Vogel & Müller-Doblies 2011).

Seed Dispersal. Myrmecochory predominates in the family (Lengyel et al. 2009, 2010).

Chemistry, Morphology, etc. The synapomorphies of tribes like Burchardieae, and of Colchicaceae as a whole, are unclear because of the uncertainty of the composition and sequence of the basal pectinations in the family. Gloriosa has leaf tendrils. Androcymbium [Colchicum] longipes has tepals ca 4.5 cm long, each with a basal claw ca 3.5 cm long representing the part of the tepal adnate to the filament.

There is considerable variation in nectary morphology and position within the family, but details of nectary evolution are unclear. Cave (1968) described Androcymbium as having a nucellar cap, although from the illustration if looks as if there is parietal tissue 2-3 cells across, similarly, illustrations in Vesque (1878) suggest that parietal tissue in Uvularia is ca 4 cells across.

Some information is taken from Nordenstam (1998); for embryology, see Cave (1968 and references), for Disporum, see Shinwari et al. (1994a, b), for Uvularia, etc., see Hayashi et al. (1998).

Phylogeny. Molecular studies suggest considerable phylogenetic structure within the family. It initially appeared that "Uvularieae" might be paraphyletic and basal. [Uvularia + Disporum - Uvularioideae A. Gray s. str.] (N. Temperate: some with disulcate pollen) and [Schelhammera + Tripladenia] (Australian) were successively the first two branches of Colchicaceae, and Drymophila was also around here (e.g. Rudall et al. 2000; Fuse & Tamura 2000; Vinnersten & Reeves 2003). These genera, and some others, have rhizomes, flavonols, and their nucellar epidermal cells are enlarged. Although Colchicaceae are noted for their alkaloids, such secondary metabolites are absent from these clades (including Uvularia: Kite et al. 2000). However, note that there is uncertainty over the relative positions of Uvularia and Burchardia (Fay et al. 2006c for a summary, see also below). Of the alkaloid-containing taxa, see the [Colchiceae [Iphigenieae + Anguillarieae]] clade above, = Colchicoideae Burmeister s. str. Note that the protoalkaloid colchicine has also been reported from some Melanthiaceae, and probably also Liliaceae and one or two other monocots not immediately related to Liliales (Tofieldiaceae, Hyacinthaceae [= Asparagaceae-Scilloideae]: Gibbs 1954), a recent survey suggests that it is restricted to Colchicaceae (Vinnersten & Larsson 2010: sampling good).

Vinnersten and Manning (2007) thought that Burchardia, sister to the rest of the family, might be paraphyletic. Androcymbium, Colchicum, Merendera and Bulbocodium form a well supported clade with (currently) little internal resolution, but the whole clade may be characterisable (Vinnersten & Reeves 2003; del Hoyo & Pedrola-Monfort 2006). For Colchicum, see Persson (2007); Androcymbium was not included there, but the recognition of Colchicum may make Androcymbium paraphyletic (see del Hoyo et al. 2009; Persson et al. 2011). The sections of Androcymbium are often not monophyletic (see also del Hoyo & Pedrola-Monfort 2008).

Classification. The tribal classification for the family that is followed here is that of Vinnersten and Manning (2007); as they note, however, there is no evidence that Burchardia (Burchardieae), sister to the rest of the family, is monophyletic; if not, an additional genus (and tribe!) may be required. Generic limits in general need much attention (Fay et al. 2006c; Vinnersten & Manning 2007). See Manning et al. (2007) for the combination of Colchicum and Androcymbium.

Previous Relationships. Genera like Drymophila were included in Luzuriagaceae by e.g. Tahtajan (1997) and placed far distant from Colchicaceae in the sequence, whilst Uvularia and Disporum used to be part of Convallariaceae (= Ruscaceae s. str., = Asparagaceae s. l.) and indeed are superficially like genera of that family such as Polygonatum, etc.

ALSTROEMERIACEAE Dumortier, nom. cons.   Back to Liliales

Leaves resupinate; (placentation parietal); testa and tegmen thin-walled; karyotype bimodal.

5/170. Central and South America, the Antipodes.

1. Alstroemerieae Bernhardi

Swollen storage roots +; flavonols, tuliposides [glucose esters] +; cuticular wax with parallel platelets; leaves spiral; inflorescence terminal, a condensed thyrse (flowers axillary), bracteoles lateral; flowers monosymmetric, T spotted, median member of the outer whorl adaxial, inner whorl often with nectariferous claw; A latrorse, basi/centrifixed; ovary inferior, stigma wet; ovules many/carpel, nucellar cap 0, micropyle?; fruit capsular [?type] or indehiscent; testa also ± thick-walled, (sarcotesta), tegmen collapses; embryo short to medium; n = 8, 9, chromosomes 6-19µm long; cotyledon not photosynthetic (photosynthetic - annual Alstroemeria graminea]).

3[list]/165: Bomarea (100), Alstroemeria (65). [Photo - Flower, Fruit.] Tropical and temperate Central and South America (map: from Aker & Healy 1990; Hofreiter 2006 - the cultivated Bomarea edulis is particularly widely distributed).

2. Luzuriageae Bentham & Hooker

Stems perennial; chelidonic acid, fructans?; leaves two-ranked, conduplicate or supervolute, petiolate or not, sheath open; pedicels articulated; T rather small; (A introrse); style deeply divided [Drymophila], stigma dry; few to many ovules/carpel, ?morphology?; fruit a berry; seeds rounded, (exotesta shed); endosperm development?; n = 10; cotyledon ?not photosynthetic, ?primary root.

2[list]/5. Peru to Tierra del Fuego, Falkland Islands, New Zealand and Australia (New South Wales to Tasmania). [Photos - Collection, Luzuriaga polyphylla, Luzuriaga radicans, Luzuriaga Flower.]

Synonymy: Luzuriagaceae Lotsy

• Alstroemeriaceae may be recognised by their leaves that are twisted at the base and so are held more or less "upside-down" and have quite broad blades.

• Alstroemerieae have in addition spirally-arranged leaves and large, monosymmetric flowers that are held with the median member of the outer whorl adaxial (and so are also "upside down"); the ovary is inferior. Luzuriageae, on the other hand, are shrubby to climbing monocots with more or less woody stems that have two-ranked leaves and undistinguished white polysymmetric flowers with plain-colored tepals and superior ovary.

Evolution. Divergence & Distribution. Stem group Alstroemerieae are dated to ca 76 million years before present, crown group Alstroemerieae to ca 30 million years before present; stem group Luzuriageae are dated to ca 79 million years before present, crown group Luzuriageae to ca 56 million years before present (Janssen & Bremer 2004).

Chemistry, Morphology, etc. Some information on Alstroemerieae is taken from E. Bayer (1998: general), Sanso and Hunziker (1998: cytology), Aagesen and Sanso (2003: phylogeny), Hofreiter (2007: biogeography and ecology), and Sarwar et al. (2010: pollen). Information on Luzuriageae is taken from Conran and Clifford (1985 [e.g. stigma], 1998).

Phylogeny. See Rudall et al. (2000a) and also Sanso and Xifreda (2001: Alstroemerieae).

MELANTHIACEAE Borkhausen, nom. cons.   Back to Liliales

Leaves often evergreen; flavones, flavonols or flavonoids +; cuticle wax with parallel platelets; (leaf margins toothed), base sheathing; inflorescence (branched) racemose; (T 3, 4); A (latrorse; adnate to base of T); G (1) [3], placentation axile to parietal, styles separate, stigma dry (wet); ovules many/carpel, position variable, parietal tissue ca 1 cell across; T persistent in fruit; seed coat?, (phlobaphene +); endosperm helobial, embryo short (large); chromosomes 1-6 µm long; cotyledon bifacial or not, hypocotyl at most short.

16[list]/170 - five groups below. N. temperate, esp. East Asia and E. North America connections, to Peru (map: see Meusel et al. 1965; Frame 1990; Fl. N. Am. 26: 2002; Seberg 2007). [Photos - Collection 1] (for Paridae, see below).

1. Melanthieae

± Bulbous; styloids also +; (leaves curved-plicate, sheath closed [Veratrum]); highly oxygenated esterified C-nor-D homosteroidal alkaloids; anthers kidney bean-shaped, opening by valves, thecae confluent; (antipodal cells persistent); capsule septicidal [ventricidal]; (seeds flat, winged); n = 8 (10, 11).

7/100: Veratrum (50), Schoenocaulon (25). North Temperate, Schoenocaulon to Peru.

Synonymy: Veratraceae Salisbury, Xerophyllaceae Takhtajan

[Helionadeae + Chionographideae]: calcium oxalate crystals cuboidal; bracts 0; anther thecae ± confluent; pollen intectate.

2. Helionadeae

Raphides 0; pollen spinulate; (style +, long); seeds linear, long-caudate at both ends; n = 17.

1/9. E. North America, East Asia.

Synonymy: Heloniadaceae J. Agardh

3. Chionographideae

Flowers often imperfect; T with 1 nerve, nectaries 0; pollen 4-porate, with clavate processes; seeds winged (at one end); n = 6?, 12 (21, 22).

2/6. E. North America, East Asia.

Synonymy: Chionographidaceae Takhtajan

[Xerophylleae + Parideae]: anther thecae distinct.

4. Xerophylleae

Plant ± bulbous; pericycle 2-3 cells thick; styloids also +; leaf long-linear, xeromorphic; T nectaries 0; ovules 2-4/carpel; n = 15.

1/2. North America.

Synonymy: Xerophyllaceae Takhtajan

5. Parideae

Rhizome monopodial; flavonols +; dimorphic root hypodermis +; raphides 0, cuboidal crystals +; stomata tetracytic; leaves whorled, with (petiole), midrib, and broad blade, venation reticulate; flowers single, terminal, (to 12-merous); P = K + C, K (0) 3-10, C (0) 3-6(-8); A 6-24, introrse to extrorse; G [3(-10)], (septal nectaries +), placentation axile to parietal, style or styluli +, stigma dry; ovules many/carpel, position variable, parietal tissue 1-2 cells across, nucellar cap 2-4 cells across, podium short, raphides +; embryo sac bisporic, 8-nucleate [Allium-type]; fruit a berry or (septicidal and) loculicidal capsule, K, C and A persistent; seeds rounded, (aril or sarcotesta +); endosperm nuclear or helobial [Trillium], embryo minute, undifferentiated; n = 5, chromosomes heteromorphic, 6-40+µm long; cotyledon unifacial (with petiole and lamina - Paris).

3-5/ca. 80: Trillium (50, or 70 when circumscribed broadly). North Temperate (for map, see Farmer 2006). [Photos: collection.]

Synonymy: Paridaceae Dumortier, Trilliaceae Chevallier, nom. cons.

Evolution. Divergence & Distribution. Melanthiaceae: stem age - ca 107 million years before present; crown age - ca 97 million years before present (Janssen & Bremer 2004).

Bacterial/Fungal Associations. Melanthieae are susceptible to infection by rust fungi (Holm 1982).

Chemistry, Morphology, etc. Narita and Takahashi (2008, see also Takahashi 1994) discuss shoot and floral development in Parideae, noting that in apetalous Trillium, but not in apetalous Paris, the carpels are alternate with the petals; they think the petals are derived from stamens, although no other Liliales have the three whorls of stamens that this hypothesis would entail (see also Ronse de Craene 2010). Although the embryo of Trillium is minute when the seed is dispersed, it grows to about the length of the seed before germination. There is no fructose oligosaccharide accumulation in Trillium, at least.

The alkaloids of Veratrum and its relatives are very complex and distinctive (Kite et al. 2000). Xerophyllum is particularly distinctive in its vegetative anatomy (Ambrose 1975). Except for Helionadeae, syncarpy in Melanthiaceae tends to be rather weak. In a number of taxa, including Veratrum and Paris, the tepals become greener and persist in fruit (e.g. Weberling 1989); other taxa, including Trillium, have persistent sepals, etc. The range of chromosome sizes in this family is at least 10-fold and that of C-values, 100-fold (Pellicer et al. 2010a and references).

For embryological information, see Berg (1962) and Cave (1968) and references. General information is taken from Ambrose (1975, 1980), Tamura (1998 - Melanthiaceae, Trilliaceae), Zomlefer (1996 - Trilliaceae, 1997a [nice table] and especially 2001 - Melanthiaceae) and Zomlefer et al. (2006); see also Arber (1925).

Phylogeny. Possible relationships are [Veratrum [Trillium and relatives + the rest] (Tamura et al. 2004a; see also Fuse & Tamura 2000), however, Xerophyllum was not included and support for the basal dichotomy was weak, or Melanthieae [[Heloniadeae + Chionographideae] [ Xerophylleae + Parideae]] (Zomlefer et al. 2006). See Farmer (2006) for detailed discussion of the Trillium group (as Trilliaceae); the backbone of the tree of relationships in Parideae remains distinctly poorly supported (see also Kazempour Osaloo & Kawano 1999 for a phylogeny).

• Parideae - Leaves pseudoverticillate, with reticulate venation; flower rather large, single, terminal; sepals and petals distinct, free; fruit berry or capsule; seed often fleshy, with sarcotesta or aril. In the past this has been recognsied as one or two separate families - Trilliaceae, sometimes also Paridaceae; generic limits are uncertain.

Previous relationships. Veratrum in particular looks superficially like Maianthemum (inc. Smilacina), a member of Ruscaceae s. str., Asparagaceae s.l.; these and most other Melanthiaceae were all included in Cronquist's (1981) Liliaceae.

[[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]: leaves broad, with reticulate venation; flowers rather small; fruit a berry.

Evolution. Divergence & Distribution. This clade diverged from other Liliales ca 115 million years before present (stem age) and started branching (crown group age - Liliaceae versus the rest) ca 91 million years before present (Janssen & Bremer 2004). Other stem and crown group ages for families in this group can be found there; note that the topology of their tree differs from that given here.

Phylogeny. It has been suggested that Philesiaceae could be included in Smilacaceae (Rudall et al. 2000) - see pollen, endosperm storage, disintegrating testa, absence of stem fructans, etc. Note, however, that in some studies (e.g. Patterson & Givnish 2002) although Philesia and Rhipogonum form a strongly associated pair, Smilax may be closer to Liliaceae, albeit only with weak support (see also introduction to the order, above). See Patterson and Givnish (2002) for characters of this group.

[Philesiaceae + Rhipogonaceae]: stem fructans 0; cuticular wax with parallel platelets; leaves with petiole, lamina, midrib; ovules crassinucellate; testa disintegrates.

PHILESIACEAE Dumortier, nom. cons.   Back to Liliales

Rhizome short and woody, plant shrub or vine, epiphytic; chelidonic acid?; velamen +; tannin and mucilage cells 0; vessels also in stem; (stomata ?type, oriented transverse to long axis of leaf [Philesia]); leaves two-ranked or spiral, conduplicate-flat or curved, with 3 main veins; (P = K + C - Philesia); A connate basally, (free), anthers dehiscing by pores, (introrse - Lapageria); pollen inaperturate, spinose; placentation intrusive parietal, stigma dry or wet; ovules many/carpel; seed exo- and endotegmic; endosperm development?, with aleurone layer and fatty oils; n = 15, 19, chromosomes 2.5-12 µm long, bimodal; cotyledon not photosynthetic, primary root well developed.

2[list]/2. S. Chile. [Lapageria Flower.]

• Philesiaceae are more or less epiphytic rather woody shrubs or vines with petiolate leaves and large, pendulous flowers; the fruit is a berry.

Chemistry, Morphology, etc. Some information is taken from Conran and Clifford (1985, 1998).

Synonymy: Lapageriaceae Kunth

RHIPOGONACEAE Conran & Clifford   Back to Liliales

Prickly climbers with short woody rhizomes; flavonols +; vessel elements in stems, with scalariform perforation plates; mucilaginous cells +; stomata unoriented; leaves opposite, conduplicate or curved, with 5 main veins; inflorescence various; T rather small; A latrorse to introrse; tapetum amoeboid; pollen prolate, reticulate; style short, stigma wet; 2 ovules/carpel; seeds rounded (subangled); exo- and endotegmen with cuticle; endosperm with starch, ?embryo; n = 15, dimorphic; seedling?, ligule 0.

1[list]/6. New Zealand to New Guinea (map: from Conran & Clifford 1986).

• Rhipogonaceae are climbers that often have prickly stems, but no tendrils. The leaves are often opposite, petiolate, and with a blade that has three, strong longitudinal veins and reticulate fine venation. The flowers are rather small and undistinguished; the fruit is a berry.

Evolution. Divergence & Distribution. Fossil Rhipogonum is reported from the Eocene of Tasmania (Conran et al. 2009b).

Chemistry, Morphology, etc. Some information is taken from Arber (1925), but see especially Conran and Clifford (1985 - e.g. vernation, seedling) and Conran (1998: general, under Smilacaceae); for pollen, see S.-C. Chen et al. (2006b).

[Smilacaceae + Liliaceae<]/a>: x = 8.

SMILACACEAE Ventenat, nom. cons.   Back to Liliales

Prickly climbers with short woody rhizomes; flavonols +, stem fructans 0; vessel elements in stem (0), with scalariform perforation plates; stem bundles in a ring; mucilaginous cells +; cuticle ± with parallel platelets, stomatal type irregular, unoriented; leaves two-ranked, conduplicate or supervolute-involute, with petiole, midrib and lamina, with 5-7 main veins, (vein endings free), paired lateral ligules and/or spines or tendrils +; plant dioecious; T (with single trace), median member of outer whorl adaxial, (± connate); staminate flowers: A 3-12, (± connate - Heterosmilax), A latrorse to introrse, bisporangiate/monothecal; nectariferous trichomes on A; pollen inaperturate, ± spheroid, ± spinulose, exine thin, endexine thick; carpellate flowers: staminodes +; style short/0, stigma dry; ovule 1/carpel, apical, straight, crassinucellate; seeds rounded (subangled), testa disintegrates, exo- and endotegmen with cuticle; endosperm with aleurone and fatty oils; embryo minute to small; n = 14-16, chromosomes 1.25-9.7 µm long; cotyledon not photosynthetic, primary root well developed, epicotyl elongated, ligule +.

1[list]/315. Pantropical to temperate (map: from Fl. N. Am. 26: 2002; FloraBase 2005). [Photo - Flower, Fruit.]

• Smilacaceae are often prickly climbers. The leaves are spirally-arranged and petiolate, there are paired petiolar tendrils, and the blade has several strong longitudinal veins and reticulate fine venation. The flowers are rather small, "typically monocot", and borne in umbels; the fruit is a berry.

Chemistry, Morphology, etc. There are vessels in the leaves (also in Eustrephus, see Laxmanniaceae, Asparagales). Smilax tends to have trilacunar nodes, and there has been some debate as to whether the paired tendrils are stipules or not (Colomb 1887; Sinnott " Bailey 1914). The blade of Smilax develops from the upper part of the leaf primordium (Bharathan 1996).

Some information is taken from Arber (1925), but see especially Conran and Clifford (1985 - e.g. vernation, seedling) and Conran (1998: general); for pollen, see S.-C. Chen et al. (2006b).

Phylogeny. Molecular analyses find the Old and New World species of the genus largely in separate clades, a result not found in morphological analyses (cf. Cameron & Fu 2006 and S.-C. Chen et al. 2006a). In the former study, Smilax aspera is sister to the rest of the family (S. vitiensis not included), in the latter, S. vitiensis is in this position and S. aspera is apparently well embedded, although not with much support (see also Kong et al. 2007 for a phylogeny and discussion on karyotype evolution).

Previous Relationships. Smilacaceae of Cronquist were broadly circumscribed and are now scattered throughout Liliales, while some are placed in Asparagales (Laxmanniaceae, see above).

LILIACEAE Jussieu, nom. cons.   Back to Liliales

Flavonols +, non-protein amino acids?, chelidonic acid 0; dimorphic root hypodermis +; raphides 0; bracteole lateral; T often spotted; A (latrose), anthers often centrifixed; (pollen operculate); (placentation parietal), stigma undivided, dry (wet); ovules 2-many/carpel, ± pendulous, (outer integument 3-6 cells across), nucellar cap (0-)1-3 cells across, (podium short), (funicular obturator +), raphides 0; antipodal cells not multinucleate; seeds flattened, stacked on top of one another, or rounded to angular; testal cells all ± thickened, some with brown contents; endosperm nuclear, embryo also short; cotyledon ± photosynthetic, bifacial, hypocotyl 0.

19[list]/610: three groups below. North Temperate, especially East Asia and North America (map: see Meusel et al. 1965; Fl. N. Am. 26: 2002).

1. Lilioideae Eaton

Plant often bulbous, with contractile roots; tuliposides, [gamma]-methyleneglutamic acid, di- and triferulic acid sucrose esters, (steroidal alkaloids) + [distribution?]; stem unbranched; leaves with parallel venation, reticulum not developed; flowers often large; style 0 or long, stigma crested; nucellar cap?, funicle large; embryo sac tetrasporic, three chalazal megaspores fuse, divide twice [Fritillaria-type]; capsule septicidal, (fruit a berry - Clintonia); seeds often flattened, (elaiosomes +); exotesta palisade or lignified, tegmen also persisting; endosperm pentaploid, thick-walled, not pitted [Erythronium]; n = 9, 11-14, [x = 12?], chromosomes (1.1 [Gagea]-)5-27 µm long; (cotyledon unifacial; not photosynthetic - Lilium canadense [Arber 1925]).

11/595: Fritillaria (130: for a classification, see Rix 2001), Lilium (110), Gagea ([70-]90[-275]), Tulipa (90). (Cold) temperate, esp. North America, East Asia. [Photos - Collection] [Photo - Nectaries]

Synonymy: Erythroniaceae Martynov, Fritillariaceae R. A. Salisbury, Liriaceae Borkhausen, Medeolaceae Takhtajan, Tulipaceae Borkhausen

[Calochortoideae + Streptopoideae]: plant rhizomatous; style divided, at least at the apex.

2. Calochortoideae Dumortier

(Bulbs); ([gamma]-methyleneglutamic acid +); (vessels in stem); leaves with parallel venation, reticulum not developed; flowers large; (K and C quite distinct), tepals pubescent; T nectaries ± saccate; capsule septicidal; n = ? Other characters to be reassigned: Placental epidermis papillate, styles ± evident; (capsule septicidal), chromosomes 1.1-6.5(-15) µm long; hypocotyl short?

5/100: Calochortus (65), Tricyrtis (20). Temperate East Asia and E. North America (map: from Fl. N. Am. 26: 2002).[Photo - Flower.]

Synonymy: Calochortaceae Dumortier, Compsoaceae Horaninow, nom. illeg., Tricyrtidaceae Takhtajan, nom. cons.

3. Streptopoideae

(P whorls very different - Scoliopus); nucellar podium very long, thin; embryo sac bisporic, 8-nucleate [Allium-type] - Streptopus; seeds striate; n = 8 (9), chromosomes 1.4-5.6(-13.2) µm long.

3/15. N. (cool) temperate, esp. East Asia and E. and W. North America (map: from Fl. N. Am. 26: 2002).

Synonymy: Scoliopaceae Takhtajan

• Liliaceae are more or less bulbous geophytes that may be recognised by their rather large flowers with six, free and often spotted tepals, six extrorse stamens, and a superior ovary.

Evolution. Divergence & Distribution. Patterson and Givnish (2002) emphasise the similarities among the large-flowered heliophilous Liliaceae, with their bulbs, capsules, and linear leaves with parallel venation, and the broad-leaved, reticulate-veined, smaller-flowered, rhizomatous, baccate, woodland Liliaceae (e.g. Prosartes, Tricyrtis) respectively, and suggest that the latter morphology is plesiomorphous in this part of Liliales ("concerted convergence" and "concerted plesiomorphy": see also Givnish 2003; especially Givnish et al. 2004b, 2005, 2006b).

Bacterial/Fungal Associations. For fungi on Liliaceae s.l., see Savile (1961).

Genes & Genomes. At least some mitochondrial genes show an accelerated rate of change in this clade (G. Petersen et al. 2006). For genome size and chromosome length, see Peruzzi et al. (2009), for chromosome size in Lilioideae, see Xie et al. (1992), and for nuclear genome size in Tulipa, see Zonneveld (2009).

Chemistry, Morphology, etc. For the distribution of tuliposide and the possibly biosynthetically related [gamma]-methyleneglutamic acid, the latter reported also from Haworthia (Asphodelaceae - Asparagales, see Fowden and Steward (1957) and Slob et al. (1975). Lilium, at least, has storage mannans in the vegetative tissues (Meier & Reid 1982).

In Streptopus (Streptopoideae) the pedicel is adnate to the stem.

Scoliopus has a differentiated perianth and only three stamens. The flowers of Lilium are shown with the median member of the outer whorl in the adaxial position (Spichiger et al. 2004; see also Eichler 1874). The pollen grains of the family are relatively large (e.g. 74-139 µm long - Handa et al. 2001), and the sulcus of Lilium can be seen even under a dissecting microscope. Medeola has been described as having crassinucellate ovules and lacking a nucellar cap, but illustrations suggest that it has an ovule rather like that of other Liliaceae (Berg 1962 and references). In Clintonia the chalazal megaspores degenerate and the endosperm is diploid (Lord 2009). There are a variety of seed dormancy mechanisms in the family, and the embryo may grow extensively after dispersal but before germination - Cardiocrinum is an example (Kondo et al. 2006).

Some general information is taken from Schnarf (1948) and Tamura (1998 - Calochortaceae, Liliaceae).

Phylogeny. The limits of the family adopted here are congruent with a phylogeny presented by Hayashi and Kawano (2000: sampling poor). Calochortus and relatives are not monophyletic in Rudall et al. (2000), but their paraphyly is not clear, either. However, support in general is stronger in Patterson and Givnish (2002) and Rønsted et al. (2005), although that for the [Calochortoideae + Streptopoideae] clade is still not very high; that support is also slight in Fay et al. (2006c: two genes, sampling poor), and the position of Tricyrtis as sister to remaining Liliaceae is also only weakly supported. In the summary tree in Peruzzi et al. (2009), Lilioideae and Streptopoideae are well supported, as is some structure within the former.

For a phylogeny of Fritillaria and Lilium, both monophyletic, see Rønsted et al. (2005a), for a more detailed phylogeny of Lilium, see Lee et al. (2011), and for that of Gagea and Lloydia, the latter para/polyphyletic and to be included in the former, see Peterson et al. (2008) and Zarrei et al. (2009). Hybridization is important in speciation in Gagea (Peterson et al. 2009).

Streptopus, Scoliopus and Prosartes make up the three genera included in Streptopoideae (see Shinwari et al. 1994a, b; Fuse & Tamura 2000; Rudall et al. 2000; and Chen et al. 2009: chemistry), etc.

Previous Relationships. Cronquist (1981) and many earlier authors circumscribed Liliaceae very broadly - Cronquist included some 280 genera and 4,000 species - and erstwhile members of the family are now scattered widely through Liliales and Asparagales in particular. Streptopus and Scoliopus have been included in a narrowly-drawn Uvulariaceae, and other members of that family are in Colchicaceae above), while Prosartes used to be included in Disporum, also now in Colchicaceae (Shinwari et al. 1994a, b).