LIGNOPHYTA

Plant a shrub or tree; true roots +, origin endogeneous, root cap +, apex multicellular; endodermis +; shoot apical meristem multicellular; lateral meristems +, cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially; lamina with mean venation density 1.8 mm/mm² (to 5 mm/mm²).

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, so no Maüle reaction; root xylem exarch, cork cambium deep seated; arbuscular mycorrhizae +; shoot apical meristem interface specific plasmodesmatal network; stem with vascular tissue around central pith [eustele], vascular bundles with interfascicular tissue, ectophloic, endodermis 0, xylem endarch; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, sieve tube plastids with starch grains; phloem fibres +; stem cork cambium superficial; branches exogenous; leaves with single trace from vascular sympodium ["nodes 1:1"]; vascular bundles collateral [stem: phloem external; leaf: phloem abaxial]; stomata morphology?, pore opening controlled by abscisic acid; leaves with petiole and lamina, spiral, development basipetal, blade simple; axillary buds +, not associated with all leaves; prophylls two, lateral; plant heterosporous, sporangia borne on sporophylls; microsporophylls aggregated in indeterminate cones/strobili; true pollen +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, parietal tissue 2+ cells across, 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, flagellae numerous; ovules increasing considerably in size between pollination and fertilization, female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large" [ca 8 mm³], but not much bigger than ovule, with morphological dormancy; embryo cellular ab initio, endoscopic, plane of first cleavage of zygote transverse, suspensor +, short-minute, embryo straight, shoot and root at opposite ends [allorrhizic], white, cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, 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, exodermis +; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, associated gelatinous fibres [g-fibres] with innermost layer of secondary cell wall rich in cellulose and poor in lignin; 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 cell and sieve tube from same mother cell; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, secondary veins pinnate, overall growth ± diffuse, venation hierarchical, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, ± haplomorphic, 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, endothecium +, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, cells 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 superior, 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]; ovule not increasing in size between pollination and fertilization; pollen binucleate at dispersal, male gametophyte trinucleate, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing between cells, growth rate 20-20,000 µm/hour, outer wall pectic, inner wall callose, with callose plugs, penetration of ovules via micropyle [porogamous], whole process takes ca 18 hours, distance to first ovule 1.1-2.1 mm; male gametes lacking cell walls, flagellae 0, double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; seed exotestal, becoming much larger than ovule at time of fertilization; endosperm diploid, cellular [micropylar and chalazal domains develop differently, 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; embryogenesis cellular; 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]].

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

[AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: essential oils in specialized cells [lamina and P ± pellucid-punctate]; tension wood 0; tectum reticulate; anther wall with outer secondary parietal cell layer dividing; carpels plicate; nucellar cap + [character lost where in eudicots?]; 12BP [4 amino acids] deletion in P1 gene.

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]] / MESANGIOSPERMAE: benzylisoquinoline alkaloids +; polyacetate derived anthraquinones + [?level]; outer epidermal walls of root elongation zone with cellulose fibrils oriented transverse to root axis; P more or less whorled, 3-merous [possible positiion]; embryo sac bipolar, 8 nucleate, antipodal cells persisting; endosperm triploid; ?germination.

[MONOCOTS [CERATOPHYLLALES + EUDICOTS]]: (veins in lamina often 7-17mm/mm² or more [mean for eudicots 8.0]); (stamens opposite [two whorls of] P); (pollen tube growth fast).

MONOCOTYLEDONS / MONOCOTYLEDONEAE / LILIANAE Takhtajan

Plant herbaceous, perennial, rhizomatous, growth sympodial; non-hydrolyzable tannins [(ent-)epicatechin-4] +, neolignans, benzylisoquinoline alkaloids 0, hemicelluloses as xylans; root apical meristem?; root epidermis developed from outer layer of cortex; trichoblast in atrichoblast [larger cell]/trichoblast cell pair further from apical meristem, in vertical files, or hypodermal cells dimorphic; endodermal cells with U-shaped thickenings; cork cambium in root [uncommon] superficial; root vascular tissue oligo- to polyarch, medullated, lateral roots arise opposite phloem poles; primary thickening meristem +; vascular bundles in stem scattered, (amphivasal), closed, vascular cambium 0; vessel elements in root with scalariform and/or simple perforations; tracheids only in stems and leaves; sieve tube plastids with cuneate protein crystals alone; stomata parallel to the long axis of the leaf, in lines, brachyparacytic; leaves with broad sheath plus blade [not petiole plus lamina], blade linear, main venation parallel, veins joining successively from the outside at the apex, endings not free, margins entire, (teeth spiny), Vorläuferspitze +, leaf base sheathing, sheath open, colleters [intravaginal squamules] +; prophyll single, adaxial; inflorescence terminal, racemose; flowers 3-merous [6-merous to the pollinator?], polysymmetric, pentacyclic; P = T, each member with three traces, median member of outer whorl abaxial, aestivation open, members of whorls alternating, similar, [pseudomonocyclic, each providing a sector for the T tube when present]; stamens = and opposite each T member [primordia often associated, and/or A vascularized from tepal trace], anther and filament more or less sharply distinguished, anthers subbasifixed, endothecium from outer secondary parietal cell layer, inner secondary parietal cell layer dividing; G [3], with congenital intercarpellary fusion, opposite outer tepals [thus median member abaxial], placentation axile; ovule with outer integument often largely dermal in origin, parietal tissue 1 cell across; antipodal cells persistent, proliferating; fruit a loculicidal capsule; seed testal; endosperm with distinct nuclear and chalazal chambers, embryo long, cylindrical, cotyledon 1, apparently terminal, plumule apparently lateral; primary root unbranched, not very well developed, "adventitious" roots numerous, hypocotyl short, (collar rhizoids +), cotyledon with a closed sheath, unifacial [hyperphyllar], both assimilating and haustorial; duplication producing monocot LOFSEP and FUL3 genes, [latter duplication of AP1/FUL gene], PHYE gene lost.

[ALISMATALES [PETROSAVIALES [[DIOSCOREALES + PANDANALES] [LILIALES [ASPARAGALES + COMMELINIDS]]]]]: ethereal oils 0; raphides + (druses 0); leaf blade vernation variants of supervolute-curved; endothecium develops directly from undivided outer secondary parietal cells; tectum reticulate with finer sculpture at the ends of the grain, endexine 0; (septal nectaries + [intercarpellary fusion postgenital]).

[PETROSAVIALES [[DIOSCOREALES + PANDANALES] [LILIALES [ASPARAGALES + COMMELINIDS]]]]: cyanogenic glycosides uncommon; starch grains simple, amylophobic; leaf blade developing basipetally from hyperphyll/hypophyll junction; epidermis with bulliform cellls [?level]; stomata anomocytic, (cuticular waxes as parallel platelets); colleters 0.

[[DIOSCOREALES + PANDANALES] [LILIALES [ASPARAGALES + COMMELINIDS]]]: nucellar cap 0; endosperm nuclear [but variation in most orders].

[LILIALES [ASPARAGALES + COMMELINIDS]]: (inflorescence branches cymose).

Evolution. Divergence & Distribution. The age of this clade is given as ca 124 m.y. by Janssen and Bremer (2004), rather older than the estimates in Bremer (2000); Magallón and Castillo (2009) suggested 135.7 and 120.1 m.y. for relaxed and constrained penalized likelihood datings respectively. However, Magallón et al. (2013) suggested an age of around 96.1 m.y. for this node.

Phylogeny. Barrett and Davis (2011) and Davis et al. (2013) found some support for a [Liliales + Asparagales] clade.

LILIALES Perleb  Main Tree, Synapomorphies.

(Plants geophytes); storage fructans, chelidonic acid, steroidal saponins +; (velamen +); (cuticular waxes as platelets transversely arranged in parallel series); leaves elliptical, (main veins seven or fewer), fine venation reticulate, (leaf base not sheathing); inflorescence terminal; T large (small), free, (spotted), tepal nectaries +; anthers extrorse; style often 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.

Note: Possible apomorphies are now being added throughout the site; they are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is because there is very considerable homoplasy, with variation within and between clades, for most characters. Furthermore, the basic information for all too many characters is very incomplete, often coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed...

Evolution. Divergence & Distribution. Crown group Liliales were dated to ca 117 m.y. by Janssen and Bremer (2004), rather older than the estimates in Bremer (2000); Magallón and Castillo (2009: relaxed and constrained penalized likelihood datings) estimated ca 125.5 and 114.4 m.y. respectively.

Genes & Genomes. Very large genomes with a C value of 35 picograms or more are found in some Melanthiaceae, Liliaceae and Alstroemeriaceae (Leitch et al. 2005).

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). Inflorescence morphology in Liliales needs attention. Three-trace tepals occur in some taxa here including Corsiaceae (Rudall and Eastman in Neyland 2002). Liliaceae and Alstroemeriaceae, at least, have distinctive radially-elongated endothecial cells (Manning & Goldblatt 1990). Glucomannan seed reserves are reported from some species of Liliaceae-Lilioideae and Colchicaceae-Colchiceae (Jakimow-Barras 1973). For chromosome size, 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, Schlittler (1953a) for inflorescences, Fukuhara and Shinwari (1994) for seed coat anatomy, Endress (1995b) for some floral development, and Conran et al. (2009b) for information on leaf and stomata.

Phylogeny. The topology of this part of the tree is based on Fay et al. (2006c), but there remain substantial areas of uncertainty. Combining Smilacaceae and Rhipogonaceae was an option in A.P.G. II (2003), however, Patterson and Givnish (2002), Fay et al. (2006c: weak support), Givnish et al. (2006: strong support), Chase et al. (2006) and J. S. Kim et al. (2013) have all found the clade [Philesiaceae + Rhipogonaceae]; Smilacaceae are strongly supported as sister to Liliaceae. Relationships suggested in the rbcL study of Janssen and Bremer (2004) are rather different, but neither Petermanniaceae nor Corsiaceae were included. J. S. Kim et al. (2012, esp. 2013) found a weakly-supported [Melanthiaceae + Petermanniaceae] clade forming a tritomy with Colchicaceae et al. and Liliaceae et al., and the position of Campynemataceae as sister to the rest of the order (Corsiaceae excluded) had no bootstrap support. See also Chase et al. (1995a), Rudall et al. (2000), Patterson and Givnish (2002), Vinnersten and Bremer (2001), and Chen et al. (2007: Bayesian analysis, support for many branches weak).

Analysis of 26S rDNA sequences suggested that Corsiaceae were polyphyletic; Arachnitis perhaps being sister to Thismia and/or Burmannia (Burmanniaceae-Dioscoreales: Neyland & Hennigan 2003; G. Petersen et al. 2006b: combined analysis). Neyland (2002a), analysing variation in 26S ribosomal DNA, thought that Corsiaceae were to be associated with Liliales. Although this position had only weak support, it is largely consistent with morphological evidence; Davis et al. (2004), Chase et al. (2006) and Petersen et al. (2012) also found Corsiaceae to be associated with Liliales. However, J. S. Kim et al. (2012, esp. 2013), using plastid sequences alone, found no support for a position in the order. These relationships need to be confirmed.

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

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

Synonymy: Campynematinae Reveal, Smilacineae Reveal - 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

Plant echlorophyllous, mycoheterotrophic; 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, 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. There are usually six extrorse stamens and the ovary is inferior.

Evolution. Pollination 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). The megaspore mother cell seems to have two cell layers above it; is this a nucellar cap? (Rübsamen 1986).

Chemistry, Morphology, etc. For information, see Rübsamen (1986: general), Neinhuis and Ibisch (1998: general), and Rudall and Eastman (2002: morphology).

CAMPYNEMATACEAE Dumortier   Back to Liliales

Rhizome short, vertical, or 0; vessels 0; fibrous leaf bases persistent; chemistry?; stomata type?; leaf lacking reticulate fine venation, base sheathing [?type]; inflorescence morphology?, axis bracteate; flowers medium-sized, T green, not spotted; A adnate to base of T, tapetum 2- or multinucleate; ovary ± inferior, style branches erect; ovules 3-many/carpel, crassinucellate; fruit a capsule or indehiscent, T enlarging, persistent; seeds angled, exotestal and endotegmic, with phlobaphene. or flattened; 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 m.y., crown group Campynemataceae to ca 73 m.y. (Janssen & Bremer 2004: Campynemataceae sister to all other Liliales).

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]]]: rhizome +, ± woody.

[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 climbing, with leaf-opposed tendrils; saponins 0; velamen +; vessels also in stem and leaf; stem with prickles; 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, outer integument 3-4 cells across, parietal tissue 2-3 cells across, nucellar cap ?2-layered; 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 (Map: from Fl. Austral. 46. 1986).

• Petermanniaceae are rhizomatous woody climbers with leaf-opposed tendrils and petiolate leaves that have reticulate venation. The smallish flowers, borne in cymose inflorescences opposite the leaves, 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 they become leaf-opposed when they are evicted by the strong growth of an axillary shoot.

Information is taken from Tomlinson and Ayensu (1969: anatomy), Björnstad (1970: embryology), Conran (1988: 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 but 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 + (0); 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]), styluli +, or style ± branched, or stigma with recurved lobes, wet or dry; ovules 2-many/carpel, ± ascending, orientation various, (unitegmic), micropyle bistomal; antipodal cells multinucleate; seeds rounded (subangled), strophiole or aril +; exotesta with thick walls (less so - Colchicum), phlobaphene +; endosperm (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; Fl. Austral. 46. 1986; 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; inflorescence umbellate, axis with laves; capsule septicidal; n = 24.

1/?. Australia.

Synonymy: Burchardiaceae Takhtajan

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

2. Uvularieae Meisner

Rhizomes +; leaves distichous, (pseudopetiole +); pollen disulcate; nucellar cap massive; (fruit a berry); endotesta enlarged; 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

Stem branching (shrub); leaves distichous, sheathing, or pseudopetiole +; inflorescence umbellate, or flowers single; (nectaries paired, stalked); 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; inflorescence ramose, or flowers single; nectary on filament bases; (antipodal cells persistent).

4. Colchiceae Reichenbach

(Plants climers), (leaf tendrils - Gloriosa), (rhizomatous); (vessels in stem - Sandersonia), (leaves whorled); (flowers monosymmetric - Gloriosa); (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

Flowers single; nectaries 0; n = 10, 11, 13, etc.

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

6. Anguillarieae D. Don

Inflorescence racemose or spicate, bracts 0; (tepals connate); (fruit septicidal); 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 m.y. before present, crown group Colchicaceae to ca 44 m.y. before present (Janssen & Bremer 2004). The Colchicum-Androcymbium clade may have diverged from the rest 13.4±1.5 m.y.a., probably in southwestrn Africa (del Hoyo et al. 2009).

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.

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 protoalkaloid colchicine has also been reported from some Melanthiaceae, probably also Liliaceae, as well as one or two other monocots not immediately related to Liliales (Tofieldiaceae, Hyacinthaceae [= Asparagaceae-Scilloideae]: Gibbs 1954), although a recent survey suggests that it is restricted to Colchicaceae (Vinnersten & Larsson 2010: sampling good). The alkaloid-containing taxa make up the [Colchiceae [Iphigenieae + Anguillarieae]] clade above, = Colchicoideae Burmeister s. str.; the tropolone alkaloids, with their remarkable seven-carbon rings, have given plant chemists headaches for a century or more.

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.

Additional information is taken from Nordenstam (1998; general); for embryology, see Ono (1929), Cave (1968 and references), for Disporum, see Shinwari et al. (1994a, b), for Uvularia, etc., see Hayashi et al. (1998), for the floral morphology of Kunthera, see Endress (1995b), and for bulbs, etc., see Buxbaum (1936) and Tillich (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), 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 basal clades (including Uvularia: Kite et al. 2000). There is uncertainty over the relative positions of Uvularia and Burchardia (Fay et al. 2006c for a summary); Vinnersten and Manning (2007) thought that Burchardia, sister to the rest of the family, might be paraphyletic, while J. S. Kim et al. (2013) placed Uvularieae basal and the three species of Burchardia they examined were in the next clade up.

Relationships in the rest of the family found by J. S. Kim et al. (2013) are consistent with those in the phylogeny above. 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 (see also del Hoyo & Pedrola-Monfort 2008; del Hoyo et al. 2009; Persson et al. 2011).

Classification. The tribal classification for the family that is followed here is that of Vinnersten and Manning (2007); if Burchardia (Burchardieae), sister to the rest of the family, is not monophyletic, an additional genus 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, since 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 (del Hoyo & Pedrola-Monfort 2008)

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; inflorescence ± cymose; (placentation parietal); testa and tegmen thin-walled; endosperm walls thick, pitted; karyotype bimodal.

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

1. Alstroemerieae Bernhardi

Swollen storage roots +; flavonols, tuliposides + [alpha-methylene-gamma-butyrolactone - glucose esters] ; cuticular wax with parallel platelets; leaves spiral; inflorescence subumbellate, cymose (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). Tropical and temperate Central and South America (map: from Aker & Healy 1990; Hofreiter 2006 - the cultivated Bomarea edulis is particularly widely distributed). [Photo - Flower, Fruit.]

2. Luzuriageae Bentham & Hooker

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

2[list]/5. Peru to Tierra del Fuego, Falkland Islands, New Zealand and southeast Australia, inc. Tasmania. (Map: from Fl. Austral. 46. 1986). [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 that 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 a superior ovary.

Evolution. Divergence & Distribution. Stem Alstroemeriaceae are dated to (116.7-)96.5, 93.4(-73.4) m.y., the crown group to (86.8-)64.2, 57.5(-37.8) m.y. (Chacón et al. 2012b, q.v. for more dates). Stem group Alstroemerieae are dated to ca 76 m.y., the crown group to ca 30 m.y.; stem group Luzuriageae are dated to ca 79 m.y., the crown group to ca 56 m.y. (Janssen & Bremer 2004) - the crown age in Chacón et al. (2012b) is (55.5-)35.9(-19.5) m.y. See Hofreiter (2007) and Chacón et al. (2012b) for the biogeography and ecology of the whole clade.

Genes & Genomes. Chacón et al. (2012a) found that there was extensive variability in the rDNA sites on the chromosomes, perhaps suggesting extensive genome rearrangements despite invariance in chromosome number.

Chemistry, Morphology, etc. Some information on Alstroemerieae is taken from E. Bayer (1998: general), Sanso and Hunziker (1998: cytology), 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), Sanso and Xifreda (2001) and Aagesen and Sanso (2003).

Classification. For generic limits in Alstroemerieae, see Sanso and Xifreda (2001).

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, style branched to the base, stigma dry (wet); ovules many/carpel, position variable, parietal tissue ca 1 cell across; T persistent in fruit, ± green; seed coat?, (phlobaphene +); endosperm helobial, embryo short (long); 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, to Peru (map: see Meusel et al. 1965; Frame 1990; Fl. N. Am. 26: 2002; Seberg 2007). [Photos - Collection 1.]

1. Melanthieae

± Bulbous; highly oxygenated esterified C-nor-D homosteroidal alkaloids; styloids also +; (leaves curved-plicate, sheath closed [Veratrum]); 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

[[Helionadeae + Chionographideae] [Xerophylleae + Parideae]]: ?

[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; T 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 unbranched), stigma dry; ovules many/carpel, position variable, parietal tissue 1-2 cells across, nucellar cap 2-4 cells across, podium short; 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 also helobial [Trillium], starchy, 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 m.y.; crown age - ca 97 m.y. (Janssen & Bremer 2004).

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

Chemistry, Morphology, etc. There is no fructose oligosaccharide accumulation in Trillium, at least. 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). There are raphides in the ovule, but nowhere else. Although the embryo of Trillium is minute when the seed is dispersed, it grows to about the length of the seed before germination.

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. 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 Ono (1929), Berg (1962) and Cave (1968) and references; for floral morphology, see Endress (1995b). General information is taken from Ambrose (1975, 1980), Tamura (1998: Melanthiaceae, Trilliaceae), Zomlefer (1996: Trilliaceae, 1997a [nice table], 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; J. S. Kim et al. 2013). Farmer (2006) discussed the relationships of the Trillium group (as Trilliaceae); the backbone of the phylogeny 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 recognised as one or two separate families - Trilliaceae, and sometimes also Paridaceae; generic limits are uncertain.

Botanical Trivia. The genome of Paris japonica, at over 150 picograms, is the largest known of any organism (Pellicer et al. 2010a).

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]]: leaf with broad lamina, with midrib, venation reticulate, petiolate, base not sheathing; fruit a berry.

Evolution. Divergence & Distribution. This clade diverged from other Liliales ca 115 m.y.a.; its crown group age is ca 91 m.y. (Janssen & Bremer 2004). Other stem and crown group ages for families in this group can be found there, but c.f. the topology of their tree.

See Patterson and Givnish (2002) for characters of this group.

Classification. Rudall et al. (2000) suggested that Philesiaceae could be included in Smilacaceae - see pollen, endosperm storage, disintegrating testa, absence of stem fructans, etc. - but this would not make sense given the phylogeny here.

[Philesiaceae + Rhipogonaceae]: stem fructans 0; cuticular wax with parallel platelets; ovules crassinucellate; testa disintegrates; chromosomes heteromorphic.

Classification. kim et al. (2013) suggest that the two families could be merged.

PHILESIACEAE Dumortier, nom. cons.   Back to Liliales

Plant shrub or vine, epiphytic; chelidonic acid?; velamen +; tannin and mucilage cells 0; vessels 0 [Carlquist]; (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, surface spinose; placentation intrusive parietal, stigma dry or wet; ovules many/carpel, parietal tissue ca 1 cell across; seed exo- and endotegmic; endosperm development?, with aleurone layer and fatty oils; n = 15, 19, chromosomes 2.5-12 µm long; 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); see Carlquist (2012a) for xylem anatomy.

Synonymy: Lapageriaceae Kunth

RHIPOGONACEAE Conran & Clifford   Back to Liliales

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

1[list]/6. New Zealand to New Guinea (map: from Fl. Austral. 46. 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: vernation, seedling, etc.; 1986: general), 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

Lianes or vines, climbing by prickles, stems monopodial; flavonols +, stem fructans 0; vessel elements in stem (0), with scalariform perforation plates; stem bundles in a ring; mucilage cells +; cuticle ± with parallel platelets, stomatal type irregular, unoriented; leaves two-ranked, vernation conduplicate or supervolute-involute, with 5-7 main veins, (vein endings free), paired lateral ligules and/or spines or tendrils; plant dioecious; flowers small, <8 mm across; T (with single trace), median member of outer whorl adaxial, valvate, (± connate); staminate flowers: A 3-12, (± connate - Heterosmilax), latrorse to introrse, bisporangiate, monothecal; nectariferous trichomes on A; pollen inaperturate, ± spherical, ± spinulose, ektexine thin, endexine thick; carpellate flowers: staminodes +; style short/0, branches stigmatiferous, long, stigma dry; ovule 1(-2)/carpel, apical, pendulous, straight, outer integument 6-10 cells across, inner integument ca 2 cells across, parietal tissue ca 5 cells across, hypostase +; seeds rounded (subangled), testa disintegrates, ± elastic, tegmen persistent, 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. Austral. 46. 1986; Fl. N. Am. 26: 2002; FloraBase 2005; Seberg 2007 - rather notional in parts). [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.

Evolution. Divergence & Distribution. Divergence within this clade may have begun at the end-Eocene ca 40 m.y.a. (Qi et al. 2012).

Chemistry, Morphology, etc. There are vessels in the leaves. 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). There is considerable variation in the leaf base, which may be more or less expanded and sheathing; the prophyll is drawn with a closed sheath by Andreata (1997). There are suggestions that the umbel is basically cymose in construction, and that plant growth may be sympodial (Andreata 1997). In general, the genus is very poorly known.

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), for ovules, see Martins et al. (2012).

Phylogeny. Molecular analyses result in the Old and New World species of the genus largely forming separate clades, a result not found in morphological analyses (c.f. 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; see also Qi et al. 2012), in the latter, S. vitiensis is in this position and S. aspera is apparently well embedded, although not with much support. Kong et al. (2007) discuss phylogeny and karyotype evolution.

Previous Relationships. Smilacaceae of Cronquist were more broadly circumscribed; the twelve genera he included are now scattered throughout Liliales and some are in Asparagales (see Asparagaceae-Lomandroideae).

LILIACEAE Jussieu, nom. cons.   Back to Liliales

Flavonols +, chelidonic acid 0; dimorphic root hypodermis +; raphides 0; bracteole lateral; T often spotted; A (latrose), anthers often centrifixed; (pollen operculate); (placentation parietal), stigma dry (wet); ovules 2-many/carpel, ± pendulous, (outer integument 3-6 cells across), nucellar cap (0-)1-3 cells across, (podium short), (funicular obturator +); antipodal cells not multinucleate; testal cells all ± thickened, some with brown contents; (embryo short); cotyledon ± photosynthetic, bifacial, hypocotyl 0.

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

1. Lilioideae Eaton

embryo sac tetrasporic, three chalazal megaspores fuse, divide twice [Fritillaria-type]; (elaiosomes +); exotesta palisade or lignified [level?].

11/595. (Cold) temperate, esp. North America, East Asia.

1a. Medeoleae Bentham

Clintonia type VAM; (leaves whorled), (with parallel venation); nucellar podium well developed; seeds rounded, elaiosome 0; exotesta palisade, outer wall thickened; embryo minute; n = 7, 14, 16.

2/6. North America, East Asia. [Photos - Collection.]

Synonymy: Medeolaceae Takhtajan

1b. Lilieae Ritgen

Plant bulbous, with contractile roots; tuliposides + [alpha-methylene-gamma-butyrolactone - glucose esters], [gamma]-methyleneglutamic acid, di- and triferulic acid sucrose esters +; leaves with parallel venation, reticulum not developed, (base sheathing - some Tulipa); flowers often large; style 0 or long, stigma crested/shortly lobed; nucellar cap?; capsule loculicidal; seeds often flattened; exotesta palisade or lignified?, tegmen also persisting; endosperm pentaploid, thick-walled, not pitted [Erythronium]; n = (9, 11-)12(-13), chromosomes (hetermorphic), (1.1 [Gagea]-)5-27 µm long; (cotyledon unifacial; not photosynthetic - Lilium canadense [Arber 1925]).

9/595: Fritillaria (130), Lilium (110), Gagea ([70-]90[-275]), Tulipa (90). (Cold) temperate, esp. North America, East Asia. [Photos - Collection, Nectaries.]

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

[Calochortoideae + Streptopoideae]: ovules with nucellar cap; placental epidermis papillate.

2. Calochortoideae Dumortier

(Bulbs - Calochortus); ([gamma]-methyleneglutamic acid +); (vessels in stem); (leaves sheathing), (with parallel venation, reticulum not developed); (outer T ± calycine, flowers 3-merous), tepals usu. ± bearded; nectaries ± saccate; (placentation parietal); capsule septicidal, seeds ± flattened; seed coat thin walled; n = 7-10, (11), 12.

2/85: 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); (A 3 - Scoliopus); nucellar podium well developed, very long, thin; embryo sac bisporic, 8-nucleate [Allium-type] - Streptopus; seeds ± rounded, striate, with phlobaphene; endotesta cells thickened on anticlinal and inner periclinal walls; 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) emphasized 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), 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. 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). For floral development, see Tzeng and Yang (2001); B-class genes are not expressed in the outer whorl of teopals 9see also Asparagaoideae). 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 (1929, 1948) and Tamura (1998: Calochortaceae, Liliaceae); for chromosomes in Lilieae, see Gao et al. (2012) and for those in Lilioideae, see Xie et al. (1992), for rootstock, etc., see Buxbaum (1938, 1958) and lillich (1998), for some chemistry, see L. Chen et al. (2009).

Phylogeny. The limits of the family adopted here are congruent with a phylogeny presented by Hayashi and Kawano (2000: sampling poor). The clade [Clintonia + Medeola] is consistently sister to the rest of Liliodeae (e.g. Patterson & Givnish 2002; Fay et al. 2006c), from which it differs somewhat morphologically. 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: esp. ndhF and combined trees) and Rønsted et al. (2005), although that for the [Calochortoideae + Streptopoideae] clade is still not very high. In Fay et al. (2006c: two genes), the positions of neither Calochortus and Tricyrtis had any support. In the summary tree in Peruzzi et al. (2009), Lilioideae and Streptopoideae are well supported, as is some structure within the former, Calochortus, but not Tricyrtis, linked with the latter. Indeed, the monophyly of Calochortoideae is questionable, Tricyrtis linking with Streptopoideae in J. S. Kim et al. (2013).

Streptopus, Scoliopus and Prosartes make up the three genera included in Streptopoideae (e.g. Shinwari et al. 1994a, b; S.-C. Chen et al. 2007), even appearing in morphological analyses in Patterson and Givnish (2002). Fay et al. (2006c) found the strongly supported relationships [Streptopus [Scoliopus + Prosartes]], c.f. the character optimizations in Patterson and Givnish (2002).

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). For a classification of Fritillaria, see Rix (2001)

Previous Relationships. Cronquist (1981) and many earlier authors circumscribed Liliaceae very broadly. Cronquist included some 280 genera and 4,000 species in the family, which is 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).