EXTANT SEED PLANTS

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 rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, 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 +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; 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, 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 duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less 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; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter 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; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable; P not sharply differentiated, outer members not enclosing the rest of the bud, smaller than inner members; A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, 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, binucleate at dispersal, trinucleate eventually, 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, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm; double fertilisation +, endosperm diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, 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 PHYA + C/PHYB + E gene pairs.

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, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families 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....

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates, axial parenchyma diffuse or diffuse-in-aggregate; 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, base not sheathing; inflorescence terminal; T large, (spotted), anthers extrorse, tepal nectaries alone, many tenuinucellate ovules/carpel, nucellar cap +, style long, stigma capitate; P deciduous; tegmen with cellular structure; endosperm with thick pitted walls, hemicellulosic; mitochondrial sdh3 gene lost. - 11 families, 67 genera, 1558 species.

Evolution. 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).

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? Very large genomes with a C value of some 350 picograms or more are found in some Melanthiaceae, Liliaceae and Alstroemeriaceae (Leitch et al. 2005).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 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 Reveal, Paridales Dumortier, Roxburghiales Wallich, Smilacales Lindley, Veratrales Dumortier - Lilianae Takhtajan, Melanthianae Doweld - Liliidae J. H. Schnaffner - Liliopsida Batsch, Liriopsida Brongniart

CORSIACEAE Beccari, nom. cons.   Back to Liliales

Echlorophyllous mycoheterotrophic herbs; chemistry?; vessels?; epicuticular wax platelets parallel, stomata unknown; 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, many ovules/carpel, integuments two cells across, nucellar cap?, funicle long, style short, (unbranched - Corsia), or stigma sessile; 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. 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?; 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, 3-many crassinucellate ovules/carpel, styluli +; 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. 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]]: rhizome +; (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

Climbers, rhizome woody; saponins 0; velamen +; vessels also in stem and leaf; stem with prickles and leaf-opposed tendrils; stomata anomocytic; 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, many ovules/carpel, micropyle?, stigma wet; 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. 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, but molecular data now separate them.

Colchicaceae + Alstroemeriaceae: ?

Colchicaceae, Alstroemeriaceae (and Luzuriagaceae, here included in the previous family) are are commonly linked, e.g. Tamura et al. (2004), Davis et al. (2004), Janssen and Bremer (2004), Fay et al. (2006c). For fructose oligosaccharide accumulation (only one record for Alstroemeriaceae s.l.), see Pollard 1982).

COLCHICACEAE de Candolle, nom. cons.   Back to Liliales

Flavones +, 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]), 2-many (unitegmic) ± ascending ovules/carpel, orientation various, micropyle bistomal, style branched or not, stigma with recurved lobes, wet or dry; 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; Hong 1993; 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.

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

2. Uvularieae Meisner

Rhizomes +; leaves distichous, (pseudopetiole +); 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.

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 +; leaves sheathing.

4. Colchiceae Reichenbach

(Leaves whorled); (tepals connate), nectaries median on tepal or on stamen base (0), (pollen polyporate); 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.

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. 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).

Myrmechory predominates in the family (Lengyel et al. 2009).

Chemistry, Morphology, etc. Particularly because of the uncertainty of the composition and sequence of the basal pectinations in the family, it is unclear both what the features of tribes like Burchardieae and the synapomorphies of Colchicaceae as a whole may be. 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 perianth adnate to the filament. There is considerable variation in nectary morphology and position within the family, and details of nectary evolution are unclear.

Some information is taken from Nordenstam (1998); 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. "Uvularieae" may be paraphyletic and basal. [Uvularia + Disporum - Uvularioideae A. Gray s. str.] (N. Temperate: some with disulcate pollen) and [Schelhammera + Tripladenia] (Australian) are successively the first two branches, and Drymophila is also around here (e.g. Rudall et al. 2000; Fuse & Tamura 2000; Vinnersten & Reeves 2003); they used to be part of Convallariaceae (= Ruscaceae s. str., = Asparagaceae s. l.) and are superficially like genera of that family such as Polygonatum, etc. 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: Wurmbaea, etc. - corms, (vessels in stems), leaf base sheathing, (nectary on filament bases) - 13 genera total (Colchicoideae Burmeister s. str). Note that the protoalkaloid colchicine is also found in some Melanthiaceae, and probably also Liliaceae and one or two other monocots not immediately related to Liliales (Tofieldiaceae, Hyacinthaceae: Gibbs 1954).

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 see Manning et al. (2007) for the combination. 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 will be required. Generic limits in general need much attention (Fay et al. 2006c; Vinnersten & Manning 2007).

Previous Relationships. Genera like Drymophila were included in Luzuriagaceae by e.g. Tahtajan (1997), placed far distant from Colchicaceae in nis sequence.

Synonymy: Bulbocodiaceae R. A. Salisbury, Burchardiaceae Takhtajan, Compsoaceae Horaninow, Merenderaceae Kral, Uvulariaceae Kunth, nom. cons.

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, many ovules/carpel, nucellar cap 0, micropyle?, stigma wet; 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), few to many [kind?] ovules/carpel, micropyle?, style deeply divided [Drymophila], stigma dry; 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.]

• 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 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. 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), and Hofreiter (2007: biogeography and ecology). 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).

Synonymy: Luzuriagaceae Lotsy

MELANTHIACEAE Batsch, 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, many crassinucellate ovules/carpel, position variable, styles separate, stigma dry (wet); 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; also styloids; (leaves curved-plicate, sheath closed [Veratrum]); highly oxygenated esterified C-nor-D homosteroidal alkaloids; anthers kidney bean-shaped, opening by valves, thecae confluent; 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, pollen 4-porate, with clavate processes, nectaries 0; 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; nectaries 0, 2-4 ovules/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, many crassinucellate ovules/carpel, position variable, embryo sac bisporic, style or styluli +, stigma dry; fruit a berry or (septicidal +) 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 Chevalier, nom. cons.

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

Chemistry, Morphology, etc. Parideae, often recognised as a distinct family (or two) in the past, may be recognised by their pseudoverticillate leaves with reticulate venation and their rather large, single, terminal flower with free sepals and petals; the fruit is a berry or capsule and the seed is often fleshy, having a sarcotesta or aril. Generic limits are uncertain. Narita and Takahashi (2008) discuss shoot and floral development, noting that in apetalous Trillium, but not in apetalous Paris, the carpels are alternate with the petals. 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.

For embryological information, see Berg (1962). General information is taken from Ambrose (1975, 1980), Tamura (1998 - Melanthiaceae, Trilliaceae), Zomlefer (1996 - Trilliaceae, 1997a 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. 2004; 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).

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. This clade diverged from other Liliales ca 115 million years before present and started branching (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; chelidonic acid?; velamen +; tannin and mucilage cells 0; vessels also in stem; (stomata anomocytic [Lapageria]; 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, introrse - Lapageria), pollen inaperturate, spinose, placentation intrusive parietal, many ovules/carpel, stigma dry or wet; 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 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 +; mucilaginous cells +; stomata unoriented; leaves opposite, conduplicate or curved, with 5 main veins; inflorescence various; T rather small; A latrorse to introrse, pollen prolate, reticulate, tapetum amoeboid, 2 ovules/carpel, style short, stigma wet; 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. Fossil Rhipogonum is reported from Tasmania (Conran et al. 2009b).

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

Smilacaceae + Liliaceae: ?

SMILACACEAE Ventenat, nom. cons.   Back to Liliales

Prickly climbers with short woody rhizomes; flavonols +, stem fructans 0; (vessels also in stem); stem bundles in a ring; mucilaginous cells +; cuticle ± with parallel platelets, stomata (anomocytic), 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 +; (T with single trace), (± connate; A 3-12, ± connate - Heterosmilax), A latrorse to introrse, bisporangiate/monothecal, pollen inaperturate, ± spheroid, ± spinulose, exine thin, endexine thick, 1 apical straight crassinucellate ovule/carpel, style short, stigma dry; seeds rounded (subangled), testa disintegrates, exo- and endotegmen with cuticle; endosperm with aleurone and fatty oils; embryo minute to small; n = 14-16, chromosomes 1.5-9.7 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). 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. ptyxis, 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), 2-many ± pendulous ovules/carpel, (outer integument ³3 cells across), stigma undivided, dry (wet); seeds flattened, stacked on top of one another, or rounded to angular; 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, embryo sac tetrasporic, three chalazal megasopores fuse, divide twice [Fritillaria-type], nucellar cap?, style 0 or long, stigma crested; capsule septicidal [Clintonia, etc.?]; seeds often flattened, (elaiosomes +); exotesta palisade or lignified, tegmen also persisting; endosperm pentaploid, thick-walled, not pitted [Erythronium]; n = 9, 11-14, 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 Batsch, Medeolaceae Takhtajan, Tulipaceae Batsch

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

2. Calochortoideae Dumortier

(Bulbs); ([gamma]-methyleneglutamic acid +); leaves with parallel venation, reticulum not developed; flowers large, tepals pubescent, saccate nectaries; capsule septicidal. Other characters to be reassigned: Placental epidermis papillate, styles ± evident; (capsule septicidal); seeds various; exo and endotegmic cuticles developed; n = 6-13, chromosomes 1.5-6.5(-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]

3. Streptopoideae

Seeds striate; n = 8.

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

Synonymy: Calochortaceae Dumortier, Scoliopaceae Takhtajan, Tricyrtidaceae Takhtajan, nom. cons.

• 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. 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).

For fungi on Liliaceae s.l., see Savile (1961).

At least some mitochondrial genes show an accelerated rate of change in this clade (G. Petersen et al. 2006).

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

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). Some general information is taken from Tamura (1998 - Calochortaceae, Liliaceae). For Streptopus and Scoliopus (ex Uvulariaceae, the others are in Colchicaceae above), and Prosartes (ex Disporum, also Colchicaceae) in particular, see also Shinwari et al. (1994a, b), Fuse and Tamura (2000), Rudall et al. (2000), etc.

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 Calochortoideae + Streptopoideae is still not very high; that support is 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.

For a phylogeny of Fritillaria and Lilium, both monophyletic, see Rønsted et al. (2005a), 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).

Previous Relationships. Cronquist (1981) circumscribed Liliaceae very broadly - some 280 genera and 4,000 species - and erstwhile members of the family as he circumscribed it are now scattered through Liliales and Asparagales in particular.