SUMMARY OF APOMORPHIES IN /APweb/
Note that there may be more detail in the characterisations, even those of the orders, on the individual order pages than appears below as possible apomorphies. This is because the ordinal pages are much more works in progress when it comes to character abstractions. These pages should always be consulted for other characters that may be apomorphies at various levels.
When scanning the summary of potential apomorphies below, remember the sometimes extreme problems one currently faces when trying to place characters on the tree. I repeat some of the points I made in the Introduction:
1. I may not have found all the information about a particular character, there may be disagreement over its interpretation (e.g. Austrobaileyales), or I have added unpublished information (e.g. Myrsinaceae and their relatives, nodal anatomy, etc.; Diapensiaceae, leaf ptyxis; Peridiscaceae, Centroplacaceae, etc.).
2. The sampling of nearly all molecular studies is incomplete (see Salisbury & Kim 2001 for problems caused by sampling). But not only is the sampling in molecular studies often less than we might wish, that of the morphological and chemical characters whose evolution we are interested in understanding is often also very poor.
3. I am fitting characters to a very conservative tree with many polychotomies, although the nodes that are utilised are for the most part strongly supported. This makes optimisation of characters, the assigment of character state change to a particular node on the tree, notably difficult (e.g. Madison & Madison 2002). In nearly all studies of the evolution of characters, the distributions of those characters are optimised on a more or less fully resolved tree, and the construction of supertrees may yield much more detailed hypotheses of relationships. Of course, some nodes on such fully resolved trees and/or supertrees may have little support, and optimisations based on such trees may carry correspondingly little conviction... To see the kind of thing that may happen as our knowledge of relationships improves, note the difference below between the possible synapomorphies given on the Dipsacales page for Dipsacales, and those that might remain if [Columelliaceae + Desfontaineaceae] were sister to Dipsacales. Even in some trees included here, parts have poor support, e.g. Ceratophyllaceae as sister to monocots, relationships within aquatic Alismatales, etc.
4. Exactly how one goes about optimising a character on a tree is critically important. Thus using either parsimony or maximum likelihood, making apparently reasonable suggestions about weighting gains over losses (or vice versa), or more complex assumptions along similar lines, or simply using ACCTRAN or DELTRAN to place the character on the tree (e.g. Donoghue & Ackerley 1996; Cunningham et al. 1998; Omland 1997, 1999; Ree & Donoghue 1999; Polly 2001; Webster & Purvis 2001; Ronquist 2004; Crisp & Cook 2005), may greatly affect the position of synapomorphies on trees, and hence our ideas of evolution. I have tried to indicate where there are particularly important uncertainties as to where characters should be placed on the tree.
5. Finally, although I have paid quite a lot of attention to the delimitation of the character "states" that make up all characterizations, I have not spent enough time on this critical operation. If we are interested in understanding evolution, then fitting the basic variation - not character states - to a tree in principle allows greater flexibility in understanding morphology in the context of local phylogenies (see also Stevens 2000; Endress 2005c). However, many states used here still owe their circumscriptions if not to their delimitation in the context of global phylogenetic analyses, at least to thinking about the variation shown by a particular character across all angiosperms. Such states often have arbitrary limits, and serve best to communicate "information"; whether they are in fact suitable for either phylogenetic analysis or understanding evolution are separate issues. So when interpreting the literature in which character states are optimised on a tree, one should bear in mind the problems surrounding the delimitation of states (e.g. Stevens 2000, 2006b). Furthermore, studies have rather unsurprisingly, perhaps, but importantly shown that dividing the one character into different sets of states may yield differing ideas of evolution of that character (e.g. Lamb Frye & Kron 2003; Hibbett 2004).
As one might expect, with increasing knowledge of development, etc., delimitation of states and characters does not necessarily become easier. Thus Buzgo et al. (2004) and Matthews and Endress (2005) show how hard it can be to delimit states or even to distinguish between e.g. prophylls and other floral structures, Peney et al. (2005) found that not all monosulcate pollen grains in monocots have the same developmental pathway, and therefore such pollen might not have the same ancestral state, while Reeves and Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in Lamiales and Solanales were different and Serna and Martin (2006) described similar problems with the development of hairs in Arabidopsis and Antirrhinum and Solanaceae.
Endress and Matthews (2006a) emphasize the importance of tendencies and developmental constraints when thinking about characters of clades within the rosids - many characters, or character combinations, in these clades occur in a rather sporadic fashion within the clade and apparently notably less frequently outside it. There are a number of other examples in the order pages here, ranging from the features enclosed by parentheses in groups above the level of family, e.g. "(cuticular waxes as aggregated rodlets)" for the commelinids to the discussion of ovular features in the asterids (see Cornales page), although I have kept few of these features in this page. One of the most striking examples is the distribution of N-fixation, restricted as it is to a monophyletic group of four clades, although it has arisen seven or more times independently within that clade (e.g. Clawson et al. 2004). More concrete examples of work that bears on the issue of tendencies include the findings that flowers of polysymmetrical Arabidopsis have genes like TCP1 that are expressed asymmetrically during early development, and TCP1 is a probable orthologue of the well-known CYC gene of Antirrhinum (Cubas et al. 2001; Costa et al. 2005; etc.). Parallelism might build on this underlying morphologically cryptic monosymmetry, even if details of the genetic mechanisms causing the monosymmetry evident in particular groups may be different (Reeves & Olmstead 2003; Cubas 2004 see above). It has also been suggested that for some secondary metabolites in particular there is aquisition of the capability to synthesise a particular metabolite; this is then switched off, but not lost - and is sometimes reacquired again (e.g. Wink 2003; Liscombe et al. 2005). Hence the rather spotty distribution of some of these metabolites when considered in the context of phylogenies. As a non-botanical example - but a rather nice one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been considerable parallelism (40-65 or more independent origins) in the evolution of eyes in metazoans. However, Pax 6 seems to be a master control gene that is involved in eye formation perhaps throughout all bilateralians (e.g. Gehring & Ikeo 1999; Erwin & Davidson 2002). When thinking of the association of camplylotopy and micropyle type, or integument thickness and vascularization of the integument (Endress & Matthews 2006a), one can imagine fairly simple developmental preconditions being operative: as Endress and Matthews (2006a) note, it is difficult to imagine vascular tissue developing in an integument only two cells thick. Changes here would be loosely correlated, if morphologically linked. However, the strongly correlated changes noted by Givnish et al. (2005) are ecologically linked but are presumably morphologically/developmentally independent. When there seem to be characters evolving more or less together, one needs to test to see if the changes are concentrated on certain branches of the tree (Maddison 1990; Maddison & Maddison 2000).
I use Remane's three main criteria of "homology", special properties, position, and intermediates, when determining the basic similarity of structures on different organisms. Even these criteria may not yield an unambiguous answer as to what a structure "is", even given a solid phylogeny. As Endress (2005c) observed, a number of features - position, function, development, shape, anatomy, histology, gene activity, and relationships to other taxa that clearly have petals - can be used to distinguish a petal (for example) from other floral structures, so if a petal does not have one of these features, is it thereby not a petal? There is rampant parallelism in seed plants, yet, as mentioned elsewhere, the terms we use to describe seed plants do not replect this; to reform terms so that they relected phylogeny/synapomorphies would cause an immense amount of grief! Given our current understandings of both phylogenies and characters, evolution of some characters in which we are interested seems to be very labile indeed (e.g. see Soltis et al. 2005b), and I have tried to be very cautious when talking about character evolution.
Finally, far too many features are still described as encompassing alternatives or a range of variation. This is perhaps an unlikely situation for apomorphies for major clades, although less so for - say - ecological preferences at the species level (Hardy 2006).
As an example of the cumulative effect of some of these problems, below I suggest two sets of possible apomorphies for Dipsacales, the first (A) is that if [Columelliaceae + Desfontaineaceae] is their sister group and includes a characterisation of that larger group, the second (B) is the one given in the Dipsacales page, which simply represents the characters common to Dipsacales and is largely innocent of any considerations of their possible outgroups.
A. [Columelliaceae + Desfontaineaceae] + Dipsacales: Group I secoiridoids +; cork deep-seated; nodes ?; pericyclic fibers 0; petiole bundles arcuate; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; flowers moderate in size, G inferior, (many ovules/carpel), style long; K persistent in fruit; exotestal cells enlarged, variously thickened and lignified.
Dipsacales: True tracheids +; nodes 3:3; buds with scales; nectaries as multicellular hairs on corolla, pollen grains trinucleate; testa vascularised, (exotestal cells palisade).
But of course even these characterisations may be strongly affected by the features of the clade sister to any [[Columelliaceae + Desfontaineaceae] Dipsacales] grouping...
B. Dipsacales: Group I secoiridoids +; true tracheids +; nodes 3:3; petiole bundles arcuate; buds with scales; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; nectaries as multicellular hairs on corolla, pollen grains trinucleate; K persistent in fruit; testa vascularised, exotestal cells enlarged (palisade), variously thickened and lignified.
Also included in the synapomorphy hierarchy are brief indications of the sizes and distributions of families and subfamilies, and usually also features by which families can be recognised. If you click on a family name that is highlighted, you will be taken to that family on the relevant ordinal page. You can get back here from any ordinal page by clicking on the highlighted "Apomorphies" immediately after the ordinal name.
SEED PLANTS
Plant woody, evergreen; true roots present; nicotinic acid metabolised to trigonelline; lignins rich in guaiacyl units; ectophloic eustele [in stem] +, endodermis 0; stem xylem endarch, root xylem exarch; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem differentiating internally, phloem externally]; wood homoxylous, tracheids +; sieve tube/cell plastids with starch grains; phloem fibers +; nodes 1:1; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral; plant heterosporous; true pollen [microspores] +, mono[ana]sulcate, pollen exine and intine homogeneous, ovules unitegmic, crassinucellate [megasporangium eusporangiate], megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development endo/exosporic, gametes two, with cell walls; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", embryo straight, endoscopic, short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication, mitochondrial nad1 intron 2 present.
Biflavonoids +; cuticle wax tubules with nonacosan-10-ol; ferulic acid ester-linked to primary unlignified cell walls; side wall pits in tracheids circular, bordered, with tori; phloem with sieve and Strasburger cells, the sieve area with pores joining to form median cavity in the region of the middle lamella; stomata haplocheilic; transfusion tissue +; microsporophylls forming determinate strobili/cones, pollen tecate, infratectum alveolate [esp. saccate pollen], endexine lamellate at maturity; ovule with pollen chamber [developing by breakdown of nucellar cells]; pollination droplet +, fertilisation 4-6 months or more after pollination, pollen tube breaks down sporophytic cells and grows away from ovule, gametophyte of tube cell, stalk cell, two prothallial cells and two multiflagellate gametes, zooidogamy; female gametophyte monosporic, with radially-elongated cells [alveoli]; testa mainly of sarcotesta and sclerotesta, ± vascularised; chromosomes of male and female gametes line up on separate but parallel spindles, proembryo with many free-nuclear divisions; gametophyte persists in seed; genome size [1C value] intermediate, 3.5-14 pg; two copies of the LEAFY gene and three of the PHY gene.
CYCADALES Dumortier
Roots and stems with contractile tissue; ß-methylamino-L-alanine and compounds producing methylazoxymethanol +, mucilage copious; association with Nostoc or Anabaena in apogeotropic coralloid roots, root hairs 0; primary thickening meristem +; wood manoxylic, large amounts of seocndary phloem persisting; reaction wood 0; tranfusion tissue +; nodes with several girdling traces; petiole vascular bundles inverted omega shape; leaf vascular bundles amphicribral; axillary buds 0; leaves large, pinnate; cataphylls +; plants dioecious; microsporangia as synangia, many abaxial microsporangia/sporophyll, dehiscing by the action of the epidermis [exothecium]; megasporophylls with terminal sterile portion; pollen tube usually unbranched, one prothallial cell, spermatogenous cells delimited by circular anticlinal wall; germination cryptocotylar, coleorhiza +; mitochondrial nad1 intron 2 present. - 2 families, 10 genera, 305 species.
CYCADACEAE Persoon - Lignins lacking syringaldehyde [Mäule reaction negative]; hairs transparent; stem polyxylic; leaflets circinate, bases persistent, midrib +, secondary vasculature diffuse; megasporophylls not forming a determinate cone, margins lobed or toothed, (1-)3-6 erect ovules/sporophyll; seeds platyspermic. - 1/105. E. Africa and Madagascar, South East Asia to New Caledonia and Tonga.
ZAMIACEAE Horaninow - Lignins with syringaldehyde [Mäule reaction positive]; hairs coloured; leaflets flat, no midrib, secondary veins regular, subparallel; megasporophylls forming a determinate cone, peltate, 2(-3) inverted ovules/sporophyll; seeds radiospermic. - 9-10/200. Scattered throughout the tropics and subtropics.
GINKGOALES + PINALES + GNETALES: wood pycnoxylic; phloem with scattered fibres alone [Cycadales?]; axillary buds +.
GINKGOALES Gorozh.
VAM present; biflavones, non-hydrolysable tannins +, lignins lacking syringaldehyde [Mäule reaction negative]; wood pycnoxylic; side wall pits in tracheids with tori; compression wood +; leaves with two traces, venation dichotomising, open; plant dioecious; 2 pendulous microsporangia/microsporophyll, dehiscing by the action of the hypodermis [endothecium], exine thin [<2 µm across]; 2(-4) terminal erect ovules/podium, with basal collar, only inner integument vascularised; pollen tube branched, spermatogenous cells delimited by circular anticlinal wall, pollen tube penetrates between sporophytic cells, sperm cells binucleate, both nuclei fuse with female gametes; seeds with sarcoexotesta, scleromesotesta, and ± degenerating endotesta; germination cryptocotylar. - 1 family, 1 genus, 1 species [all rather redundant!].
GINKGOACEAE Engler - Plant resinous, mucilage +; short shoots +, wood there manoxylic; sclereids +; leaves deciduous. - 1/1. E. China.
PINALES + GNETALES: pollen tube unbranched, growing towards the ovule, gametes non-motile, released from the distal end of the tube, siphonogamy; germination epigeal.
PINALES Dumortier
Plants with resin; wood pycnoxylic, primary xylem endarch, bordered pits on tracheids round, opposite; side wall pits in tracheids with tori; leaves with single vein; exine thick [>2 µm across], granular; microsporangium dehiscing by the action of the hypodermis [endothecium]; ovulate strobilus compound, with ± united flattened ovuliferous and bract scales, pollen chamber 0; seed coat dry, not vascularised; proembryo with 4 or 5 nuclear divisions, with upper tier or tiers of cells from which secondary suspensor develops, elongated primary suspensor cells and basal embryonal cells [or some variant]; germination epigeal; plastid transmission paternal [?level], mitochondrial nadI gene intron 2 present. - 8 families, 68 genera, 545 species.
PINACEAE F. Rudolphi - Biflavonoids 0; sieve cells with nacreous walls, plastids with protein fibers; phloem fibers 0; 2 microsporangia/microsporophyll, pollen saccate, exine thin [<2 µm] except distally, alveolate, ovules 2/scale, inverted, pollination droplet 0 (+); sperm cells binucleate; "embryo tetrad" present [free-nuclear stage with only four nuclei]; seeds winged, wing develops from adaxial side of scale; cotyledons (2-)4-11(-20); only one copy of the chloroplast inverted repeat. - 11/210. North Temperate.
[Araucariaceae [Phyllocladaceae + Podocarpaceae]] [Sciadopityaceae [Cupressaceae + Taxaceae]]: phloem stratified; mitochondrial nadI gene intron 2 lost.
Araucariaceae [Phyllocladaceae + Podocarpaceae]: roots with nodules; one ovule/scale; proembryo with 5 or 6 free-nuclear divisions; 2nd intron in nad1 lost.
ARAUCARIACEAE Henkel & W. Hochst. - Stem apex with tunica/corpus construction; wood lacks resin canals or cells, only plugs present; pits on radial walls of tracheids touching, hexagonal in outline; stomata tetracytic; leaves multiveined; pollination droplet 0, pollen intectate, granulate; nucellus protrudes from the micropyle [?Araucaria]; pollen grains intectate, granulate; pollen germinates on ovuliferous scale and tubes grow over the scales, prothallial cells numerous; seed winged [wing the entire bract scale] or not; free nuclear stage in proembryo many nucleate, central, embryonal cells surrounded by cap cells that degenerate. - 3/33. Southern South America, Malesia to E. Australia and New Zealand).
Phyllocladaceae + Podocarpaceae: sclereids numerous, with large lumen; microsporophylls with two sporangia; pollination droplet +; male gametophytes with 3-6(-8) prothallial cells, sperm cell binucleate, whether or not one nucleus is extruded; proembryo cells binucleate[?].
PHYLLOCLADACEAE Bessey - Phylloclades +, leaves reduced to scales; pollen without sacci, pollination droplet actively resorbed; seed arillate. - 1/ca 5. The Philippines (N. Luzon) to Australia (Tasmania) and New Zealand.
PODOCARPACEAE Endlicher - Transfusion tissue in patches lateral to vascular bundles in leaf; pollen saccate, ovule with epimatium. - 17/125. Largely southern Hemisphere, scattered.
Sciadopityaceae [Cupressaceae + Taxaceae]: pollen without sacci, exine shed on germination [microgametophyte naked], prothallial cells 0.
SCIADOPITYACEAE Luersson - Leaves reduced to brown scales, short shoots with photosynthetic cladodes; microsporophyll with flattened apical expansion; 7-9 ovules/ovuliferous scale. - 1/1. C. and S. Japan.
Cupressaceae + Taxaceae: cone scales opposite.
CUPRESSACEAE Bartling - 29/140. Northern hemisphere, more scattered in south temperate region, also NE Africa.
TAXACEAE Berchtold & J. Presl - Plant dioecious; pollen inaperturate; ovule solitary, on shoot in axils of vegetative leaves; male gametes unequal in size. - 6/30. Scattered in the Northern Hemisphere, esp. South East Asia, also New Caledonia.
GNETALES Luersson
Lignins with syringaldehyde [Mäule reaction positive]; stem apex with tunica/corpus construction; roots diarch; vessels + [from circular bordered pits], both fiber tracheids and tracheids +; side wall pits in vessels with tori; mucilage cells +; leaves with two traces; stomata syndetochelic; leaves opposite, joined at the base, with collateral buds; strobilus compound, bracts opposite; plant dioecious, but micro- and sterile megasporangium-bearing structures at least sometimes closely associated; microsporangial strobilus compound, microsporangia in synangia surrounded by a tubular "bract", dehiscing apically by the action of the epidermis [exothecium], pollen striate, with granular layer under the tectum [the infratectum, together making up the ektexine]; ovules terminal, integument with much-elongated beak, surrounded by connate structure ["outer integument"]; sperm cell binucleate sperm, both gametes fuse with female gametes; secondary suspensor developing from upper embryonal tier, no primary suspensor; 5' end of the inverted repeat extended. - 3 families, 3 genera, 96 species.
EPHEDRACEAE Dumortier - Xeromorphic; nodes 1:2; leaves reduced, or at least without a lamina; microsporangiophores with 2-8 synangia, each with 2(-4) sporangia, dehiscence porose, pollen lacking a colpus, exine shed on germination; archegonia exposed at base of deep pollen chamber; each nucleus of free-nuclear stage forms an embryo; seed with papillae on the inner side of the outer covering; n = 7. - 1/65. North (warm) temperate, W. South America; drier habitats.
Gnetaceae + Welwitschiaceae: torus:margo pits 0; branched sclereids +; stomata mesogenous; male gametophyte with one prothallial cell but no sterile cell; megaspores tetrasporic, no archegonia per se, alveolation does not occur; some cells of embryonal mass elongate, embryo cellular, with lateral "feeder" [protrusion of the hypocotylar axis].
GNETACEAE Lindley - Vessels with vestured pits; sieve tubes with companion cells [derived from different cells]; laticifers +; leaves with 5 or more traces, more than two orders of reticulate venation; ovules and microsporangiophores at same node in staminate plant; microsporangiophore with (1-)2(-4) sporangia, pollen not striate, surface spinose, ovule surrounded by additional connate "integument"; elongated suspensor tubes initially formed, nucleus at end divides forming a embryonal mass; n = 11; one copy of the LEAFY gene. - 1/30. Tropical, rather disjunct.
WELWITSCHIACEAE Caruel - Successive cambia + [in root - derived from phelloderm]; leaves amphistomatic; three pairs of leaves only, the second pair persisting for the life of the plant and elongating from the base, venation parallel; ovules and microsporangiophores in intimate association, microsporangiophores 6, basally connate, with synangia of three sporangia, dehiscence radial; ovule with additional pair of bracts; megagametophyte with multinucleate cells, some grow upwards through nucellus forming female gametophytic tubes, fertilisation in apical bulge [both gametes involved?], proembryo pushed back down tube by elongating embryonal suspensor; n = 24; nad1 intron 2 absent. - 1/1. S.W. Africa.
MAGNOLIOPHYTA
Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway, 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; cork cambium in root deep-seated, trichoblasts [differentiated hair-forming cells] 0, origin of epidermis with no clear pattern [probably from inner layer of cap]; stem with 2-layered tunica-corpus construction; cork cambium in stem superficial; wood fibers and wood parenchyma +; circular bordered pits lacking margo and torus; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with sieve plate, companion cells from same mother cell that gave rise to the sieve tube and P proteins, plastids with starch grains alone; nodes unilacunar; stomata paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, A many, development centripetal, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, thecae dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther, tapetum glandular, binucleate, microsporogenesis successive, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous, endexine compact, lamellate only in the apertural regions, pollen tube elongated, with callose plug, growth moderately fast, siphonogamy, gametes 2, with cell walls, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, ovules marginal, anatropous, bitegmic, micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin, chalazal, female gametophyte ?type, stylodium short, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; 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; germination hypogeal, sympodial seedlings/young plants; 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/PHYC gene pairs.
Possible angiosperm synapomorphies are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, partly because many taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied and the immediate outgroup to angiosperms is also uncertain.
AMBORELLALES Melikian, A. V. Bobrov & Zaytzeva
Nodes 1:1; plant dioecious; hypanthium +, A sessile, vascular bundle branched towards apex, microsporogenesis successive, pollen ektexine granulate, 1 ovule/carpel, embryo sac monosporic, spore chalazal, 8-nucleate, bipolar; fruit of drupelets; endosperm triploid. - 1 family, 1 genus, 1 species.
AMBORELLACEAE Pichon - 1/1. New Caledonia.
NYMPHAEALES + AUSTROBAILEYALES + [MONOCOTS + CERATOPHYLLALES] + [CHLORANTHALES + MAGNOLIIDS + EUDICOTS]: vessels +, elements with scalariform perforation plates; pollen tectate-columellate, tectum reticulate [perforated]; nucleus of egg cell sister to one of the polar nuclei; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.
NYMPHAEALES Dumortier
Aquatic herbs; hydrolyzable [ellagi]tannins +; root epidermis with alternating long and short cells; cambium 0; 4-celled uniseriate secretory trichomes with a large terminal cell; starch grains compound; primary root soon aborts, root apex with secondary dermatogen, etc., trichoblasts in vertical files, proximal cell smaller, epidermis from outer layer of cortex, diaphragms in root aerenchyma; primary stem with ± scattered closed bundles; secondary thickening 0; nodes?; aerenchyma common; stomata anomocytic; leaf base broad; seeds exotestal, operculate; endosperm scanty, perisperm precocious, ± multinucleate, copious [starchy], suspensor 0, embryo broad; seedling/young plant monopodial; intergenic inversion in chloroplast inverted repeat. - 3 families, 7 genera, 74 species.
HYDATELLACEAE U. Hamann - Plant rosette-forming; sieve tube plastids with cuneate proteinaceous inclusions; stomata anomocytic; leaves linear, with a single vein; plant monoecious; inflorescence scapose, with involucral bracts; flowers imperfect, ebracteate; P 0; staminate flowers: A 1; carpellate flowers: G with carpellary bundles equidistant, ovule single, pendulous, tenuinucellate, stigma penicillate; perisperm precocious; endosperm cellular. - 2/10. India, New Zealand and Australia.
Cabombaceae + Nymphaeaceae: stem rhizomatous; leaves involute, peltate, 2ndary veins palmate, actinodromous, festoon brochidodromous; flowers single along stem, with cortical vascular system, P whorled (outer [inner] whorls in 3's), outer members enclosing the rest of the bud, A whorled, placentation ± laminar, carpel margins with postgenital fusion; exotesta palisade, anticlinal walls sinuous.
CABOMBACEAE A. Richard - Floating and rhizomatous; stem vascular tissue with two pairs of bundles; stomata anomocytic; flowers 3-merous, A with slender filaments. - 2/6. World-wide.
NYMPHAEACEAE Salisbury - 3/58. World-wide.
1. Nupharoideae Ito - Rhizomes stout, creeping; roots with well-developed pith; K 5-14, pollen spiny. - 1/ca 10. North Temperate.
2. Nymphaeoideae - Inner satellite peduncle bundle +; K 4-5, staminodes showy, pollen zonasulcate, G more or less inferior, stigmatic surface continuous; fruit maturation underwater; seeds arillate. - 2/48. World-wide.
AUSTROBAILEYALES + CERATOPHYLLALES + CHLORANTHALES + MAGNOLIIDS [MONOCOTS + EUDICOTS]: ethereal oils in spherical idioblasts [lamina and P ± pellucid-punctate]; tension wood 0; 12BP [4 amino acids] deletion in P1 gene.
AUSTROBAILEYALES Reveal
Tiglic acid +; nodes 1:2; P not obviously in 3's, extragynoecial mucilaginous compitum +, outer integument 5-7 cells thick; fruit a berrylet; mesotestal cells ± sclerotic. - 3 families, 5 genera, 100 species.
AUSTROBAILEYACEAE Croizat - Flowers with internal staminodes. - 1/2. Australia.
Schisandraceae + Trimeniaceae: ?
SCHISANDRACEAE Blume - Tetracyclic triterpenes +; astrosclereids +; mucilage cells +; leaf epidermis silicified; pollen tricolpate, syncolpate pole distal, semitectate-reticulate, muri tall; exotesta palisade. - 3/92. Sri Lanka and South East Asia to W. Malesia, S.E. U.S.A., E. Mexico, Greater Antilles.
TRIMENIACEAE Gibbs - Rays 6-9-seriate; flowers small, carpels 1-2, 1 ovule/carpel, micropyle bistomal. - 1-2/6. New Guinea and S.E. Australia to Fiji.
[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] : benzylisoquinoline alkaloids +; P more or less whorled, 3-merous [possible position], A whorled, carpels plicate, fusion by congenital occlusion; embryo sac bipolar, 8 nucleate; endosperm triploid.
CHLORANTHALES + MAGNOLIIDS: Sesquiterpenes +.
CHLORANTHALES J. F. Leroy
?Neolignans +; rays 6-10-seriate; nodes often swollen, ?type; leaves opposite, toothed, teeth with lateral vein and others; stipules +; flowers very small, monosymmetric, parts whorled; P 3 or 0, A 1, G 1, ± inferior, ascidiate, postgenital fusion by secretion, 1 apical atropous ovule/carpel; fruit fleshy; endotesta palisade, lignified, crystalliferous. - 1 family, 4 genera, 75 species.
CHLORANTHACEAE Sims - 4/75. Tropics and subtropics, not Africa (Madagascar - Chloranthus only).
MAGNOLIIDS: (neolignans +); leaf margins entire; A many, spiral [possible position here], extrorse, antipodal cells ephemeral, hypostase +, nucellar cap +, raphal bundle branches at the chalaza.
MAGNOLIALES + LAURALES: cuticle waxes as annularly-ridged rodlets, palmitol the main wax.
MAGNOLIALES Bromhead
Vessels in multiples; secondary phloem stratified; pith septate [with sclerenchymatous diaphragms]; nodes 3:3; petiole vasculature an arc with an adaxial plate; leaves 2-ranked; G occluded by fusion and secretion, outer integument 5-10 cells across, obturator +; seeds medium-sized, testa vascularised; endosperm ?type, irregularly ruminate. - 5 families, 154 genera, 2929 species.
MYRISTICACEAE R. Brown - Sieve elements with nuclear non-dispersive protein bodies; tannin-containing tubes in the xylem; hairs branched or stellate (T-shaped); flowers imperfect, P 3, connate; staminate flowers: A whorled, connate, unithecate, pollen aperture membrane sculpted; carpellate flowers: G 1, one ovule/carpel, inner integument 3-10 cells across; fruit a follicle also dehiscing abaxially; seed large, pachychalazal, arillate, endotesta palisade, lignified, crystalliferous, tegmen vascularised, massive, exotegmen sclerotic or tracheidal; endosperm nuclear; hypocotyl not developed. - 20/475. Pantropical.
Magnoliaceae + Himantandraceae + Degeneriaceae + Eupomatiaceae + Annonaceae: primary stem with distinct bundles [eustele]; wood with broad rays; flowers solitary, large, receptacle well-developed, cortical vascular system +, P = K + C, A many, spiral [another position!], filaments with three veins, anther thecae separate, embedded in the broad filaments, the connective prolonged, funicular obturator +; testa multiplicative; 10-aa deletion in PI-derived motif in AP3 gene.
MAGNOLIACEAE Jussieu - Sesquiterpene lactones +; nodes 6(+):6(+); stipules surrounding stem; flowers terminal; receptacle elongated, micropyle bistomal. - 2/227. The Americas (but not W. North America), and South East Asia to Malesia.
Himantandraceae + Degeneriaceae + Eupomatiaceae + Annonaceae: anthers valvate, staminodes internal, pollen with granular infratectum, atectate, psilate, pollen tube transmission tissue differentiated; fruit indehiscent.
Degeneriaceae + Himantandraceae: flowers axillary; outer integument annular; x = 12.
DEGENERIACEAE I. W. Bailey & A. C. Smith - Leaves spiral, G 1; seeds with sarcotesta. - 1/2. Fiji.
ngle carpel.HIMANTANDRACEAE Diels - Plant with peltate scales; P or bract + bracteoles enclosing the flower in 2 series, caducous, pollen scabrate, (1) 2 apical ovules/carpel; fruit ± syncarpous, a drupe with several stones; seeds flattened. - 1/2. New Guinea and N.E. Australia.
Eupomatiaceae + Annonaceae: sieve tube plastids with protein crystalloids and starch; rays 8-15-seriate; petiole bundles arcuate; trunk leaves spiral; inflorescence +; fruit ± berry-like; mesotesta fibrous.
EUPOMATIACEAE Endlicher - Inflorescence fasciculate; receptacle concave, calyptra +, thick, deciduous, with sclereids, P 0, A introrse, basally connate, pollen with encircling equatorial sulcus. - 1/3. New Guinea and E. Australia.
ANNONACEAE Jussieu - Flowers with open development; A whorled, filaments with a single vein; endotestal plug +; tegmen crushed, ruminations irregular. - 129/2220. Largely tropical.
Anaxagorea - Uniseriate hairs terminate in a rounded cell; trunk leaves distichous; receptacular vascular system 0; fruit a follicle; endotesta aerenchymatous, tegmen alone involved in ruminations, with idioblastic oil globules. - 1/21. Tropical America, also Sri Lanka to West Malesia.
Ambavia clade + The Rest: internal staminodes 0.
2. The Ambavia clade - Anther connective ± tongue-like, ovules (1)2/carpel, integuments 3. - 6/54. Tropical, inc. Madagascar.
The rest - Stamens with prominent ± truncate connective. - 128/2200. Predominantly lowland tropics.
LAURALES Perleb
Sesquiterpenes 0; sieve tube plastids with protein crystalloids; nodes 1:?; petiole bundle(s) arcuate; leaves opposite; inflorescence ± cymose; hypanthium +, inner staminodia +, ovules basal, archesporium multicellular, stylodium long; fruitlets 1-seeded, indehiscent, hypanthium persistent; mesotesta crushed, endotesta tracheidal [seed endotestal], tegmen crushed; embryo long; duplication of the PI gene. - 7 families, 91 genera, 2858 species.
CALYCANTHACEAE Lindley - Stem with four inverted cortical bundles; P many, spiral, no double fertilisation [endosperm develops autonomously]. - 5/11. China, North America, N.E. Australia.
1. Idiospermum - G 1-2(-3), outer integument 12-15 cells thick, micropyle bistomal, nucellar beak +; cotyledons (3-)4, massive, peltate. - 1/1. N.E. Australia.
2. The rest - Pollen euatorially and vertically disulcate, G 5-35, outer integument 5-6(-8) cells thick; cotyledons spirally twisted. - 4/10. China, North America.
Siparunaceae + Atherospermataceae + Gomortegaceae + Monimiaceae + Hernandiaceae + Lauraceae: vessel elements with scalariform perforations; hippocrepiform sclereids in pericycle; lamina with rather distant teeth, one vein entering opaque, persistent glandular cap; flowers rather small, A whorled, stamens with paired nectaries/glands at base, anthers bisporangiate/unithecal, valvate, tapetum ?, pollen inaperturate, ± spinulose, 1 ovule/carpel.
Siparunaceae + Gomortegaceae + Atherospermataceae: acicular crystals +; hypanthium closed by roof, infratectum columellar; embryo very small.
SIPARUNACEAE Schodde - A bisporangiate/unithecal, with a single slit, basal glands 0, integument 0. - 2/75. Tropical America, W. Africa.
Gomortegaceae + Atherospermataceae: bud scales +; sieve tube plastids also with fibrils; outer, not inner, A staminodial.
GOMORTEGACEAE Reiche - Lamina entire; G [2-3(-5)], inferior, ovule apical, stigmatic branches erect; fruit with a single seed, stone wall thick; embryo large, endosperm slight. - 1/1. C. Chile.
ATHEROSPERMATACEAE R. Brown - Stomata anomocytic; fruit achenial, plumose, hypanthium woody. - 6-7/16. New Guinea to New Zealand and New Caledonia, Chile.
Monimiaceae + Hernandiaceae + Lauraceae: A whorled, columellae foot layer and endexine 0, ovule apical.
MONIMIACEAE Jussieu - Paired nectaries/glands at base of A absent, anthers dehiscing longitudinally. - 22/200. Tropical, but esp. Australasia.
Hernandiaceae + Lauraceae: primary stem ± with vascular cylinder; vessel elements with simple perforations; mucilage cells +; sieve tube plastids also with starch; leaves spiral, entire (lobed); flower parts whorled, tapetum amoeboid, exine thin, intine very thick, G 1, ovule pachychalazal, outer integument 4< cells across, embryo sac more or less linear, hypostase 0; testa multiplicative, thick, tegmen not persisting; endosperm 0.
HERNANDIACEAE Berchtold & J. Presl - G inferior [one position]; fruit dry. - 5/55. Pantropical.
Hernandioideae - anther valves laterally hinged, tapetal cells radially elongated, single layer of microspore mother cells, pollen grains 90-160 µm across, outer integument 10-23 cells thick, nucellus massive, 6-8 layers of parietal cells, nucellar beak +; testa vascularised, spongy, mesotesta massive, 7-17 cells across. - 3/44. Tropical, esp. Madagascar and Indo-Malesia.
Gyrocarpoideae Pax - Strong higher-order vein areolation; inflorescence ebracteate; apical part of embryo sac protruding; cotyledons contortuplicate [much folded!]. - 2/10. Pantropical, esp. America.
LAURACEAE Jussieu - Leaves entire, strong higher-order vein areolation; pollen mother cells in a single row, archesporium single-celled; fruit a drupe. - Ca. 50/2500. Pantropical (temperate), lowland to montane.
1. Hypodaphnis - Anthers tetrasporangiate, staminodes 0, G inferior. - 1/1. Tropical West Africa.
Beilschmeidia, Cryptocarya, Endiandra, etc. [Cassytha [Caryodaphnopsis and Neocinnamomum + The Rest]]: subsidiary cells of paracytic stomata envelop the guard cell above and below, the latter having outer and inner cuticular ledges; staminodes +, glandular tapetum +, protruding embryo sacs.
2. Beilschmeidia, Cryptocarya, Endiandra, etc. - Stamens in two whorls - 6/710. Pantropical, some subtropical, to New Zealand.
Cassytha [Caryodaphnopsis and Neocinnamomum + The Rest]: ?
3. Cassytha - Parasitic herb; micropyle bistomal, nucellar cap 0; endosperm cellular. - 1/16. Old World tropics.
Caryodaphnopsis and Neocinnamomum + The Rest: ?
Caryodaphnopsis + Neocinnamomum - Anthers tetrasporangiate. - 2/13. South East Asia to the Philippines and Borneo.
5. The Rest - Tapetum amoeboid, embryo sac not protruding. - >Ca 40/1730. Pantropical (temperate).
CANELLALES + PIPERALES: aporphine alkaloids, flavonols +; nodes 3:3; G whorled.
CANELLALES Cronquist
Indumentum 0; sieve tube plastids with starch and protein crystalloids and/or fibres; petiole bundle(s) arcuate; leaf waxes as tubules with nonacosan-10-ol the main wax; foliar sclereids +, branched; K and C distinct, micropyle bistomal. - 2 families, 10 genera, 93 species.
CANELLACEAE Martius - Cortical vascular system +, G [2-6], also occluded by secretion, placentation parietal, ovules campylotropous, micropyle zig-zag; fruit a berry, K persistent. - 5/13. Tropics; U.S.A. (S. Florida), Antilles, South America, Africa, Madagascar.
WINTERACEAE Lindley - Vessels 0; rays 10+-seriate; K calyptrate, pollen in tetrads, monoporate. - 4-7/60-90. Montane tropics, not mainland Africa.
Taktajanioideae Leroy - Stomata anomocytic; G [2], placentation parietal. - 1/1. Madagascar.
Winteroideae - Sesquiterpene dialdehyde cinnamates + - 3-6/60-90. New Guinea to New Zealand and New Caledonia, few Borneo and the Philippines and South America.
PIPERALES Dumortier
Herbs; vessel elements with simple perforations; starch grains compound; primary stem with distinct bundles; vessel elements with simple perforations; wood with broad rays; nodes often swollen; stomata ?; leaves 2-ranked, 2ndary veins palmate; flower parts whorled, P and A in 3's, G occlusion?; seed ± tegmic. - 4 families, 17 genera, 4090 species.
Hydnoraceae + Aristolochiaceae: P connate, valvate, A extrorse, embryo undifferentiated.
HYDNORACEAE C. Agardh - Echlorophyllous root parasites; leaves 0; flowers arising endogenously from roots, large, P uniseriate, thick and fleshy, A = adnate to and opposite P, A sessile, connate, polythecate, ektexine homogeneous, G inferior, placentation of parietal or apical lamellae, many ovules atropous, unitegmic, tenuinucellate, stigma broad, cushion-shaped. - 2/7. Arabian Peninsula, Africa, Madagascar; Costa Rica and S. South America.
ARISTOLOCHIACEAE Jussieu - Stomata anomocytic; leaves conduplicate, heart-shaped, 2ndary venation palmate, prophyll adaxial; inflorescence cymose: P 3, odd member adaxial, A in 3's, ± sessile, connective extended apically, carpels basically free; fruit a follicle; endotesta palisade, usu. crystalliferous, exotegmen and layer underneath crossing fibers, endotegmen with reticulate thickenings. - 5-8/480. World-wide, not Arctic - three groups below.
1. Asaroideae O. C. Schmidt - Growth sympodial; sieve tube plastids with cuneate protein crystalloids and a large polygonal crystal. - 2/75. N. Temperate, esp. East Asia.
Lactoris + Aristolochioideae: ?
2. Lactoris - Sieve tube plastids with starch grains; nodes 1:2; leaves elliptic, 2ndary veins subpinnate, stipule sheathing, intrapetiolar, adnate to the petiole; bracteoles 0; flowers small, A 6, pollen in tetrads, saccate, G 3, ovules ?tenuinucellate, funicle long, endothelium +. - 1/1: Lactoris fernandeziana. Chile, the Juan Fernandez Islands.
See Carlquist (1964: general, 1990: wood anatomy), Crawford et al. (1986: chemistry), Metcalfe (1987: vegetative anatomy), Kubitzki (1993: general), Tobe et al. (1993) and González and Rudall (2001: general, the stipule is initially paired) for details.
3. Aristolochioideae - Growth monopodial; benzylisoquinoline alkaloids +; sieve tube plastids with polygonal protein crystalloids plus starch grains or protein fibers (starch grains alone); serially arranged axillary buds; hairs hooked; inflorescences axillary, bracts distinct; floral primordia monosymmetric, G [4-6], inferior, apically constricted; fruit septicidal and opening laterally, a schizocarp, or dry-baccate. - 2-5/405. Tropics (temperate), relatively less diverse in Africa (inc. Madagascar), few in N. Australia.
Piperaceae + Saururaceae: root epidermis from inner layer of cap; stomata tetracytic; leaf base broad, ± sheathing; inflorescence spicate, terminal; flowers small, P 0, filaments rather slender, pollen grains <20 µm, ovules atropous, stigma dry; seed coat exo- and endotegmic; perisperm +, endosperm ?type, scanty, embryo broad.
PIPERACEAE Martynov - Vascular bundles not in a single ring; flowers very small, bracts peltate; fruit a drupe. - 5/3600. Pantropical.
SAURURACEAE Martynov - Inflorescence bracts petaloid; A introrse, carpels connate, placentation parietal. - 4/6. North Temperate.
MONOCOTS [CERATOPHYLLALES + EUDICOTS]: (stamens opposite [two whorls of] P).
MONOCOTYLEDONS: Herbaceous, plant sympodial; non-hydrolyzable tannins [(ent-)epicatechin-4] +, benzylisoquinoline alkaloids, ellagitannins, neolignans 0, hemicelluloses as xylans; root apical meristem?; root epidermis developed from outer layer of cortex; trichoblasts in vertical files with proximal cell smaller or hypodermal cells dimorphic; cork cambium in root [uncommon] superficial; root vascular tissue oligo- to polyarch, medullated, lateral roots arise opposite phloem poles; tunica 2-layered [?sampling]; primary thickening meristem +; vascular bundles in stem scattered, (amphivasal), closed [no interfascicular cambium developing]; vessels in root with scalariform and/or simple perforations; vessels in stems and leaves 0; sieve tube plastids with cuneate protein crystals alone; epidermis with bulliform cells [?this level], stomata paracytic [cell divisions oblique]; leaves not differentiated into petiole plus lamina, main venation parallel, developing both acropetally and basipetally from the base and converging towards the apex, intermediate [and other] veins basipetal from apex, endings not free, (margins with spiny teeth), Vorläuferspitze +, base sheathing, sheath open, colleters [intravaginal squamules] +; inflorescence racemose; flowers 3-merous, polysymmetric, pentacyclic, T (each with three traces), in two whorls, median member of outer whorl abaxial, members of whorls alternating, similar, [pseudomonocyclic, each providing a sector for the T tube when present], A = and opposite each T member (primordia associated), anther and filament sharply distinguished, microsporogenesis successive, G [3], development?, opposite outer tepals [thus median member abaxial], placentation axile, style hollow, short; fruit a loculicidal capsule; seed testal; embryo long, cylindrical, cotyledon 1, terminal, plumule lateral; seedling phanerocotylar, primary root unbranched, adventitious roots numerous, hypocotyl short, (collar rhizoids +), cotyledon with a closed sheath, unifacial, assimilating and haustorial; duplication producing monocot LOFSEP gene.
Some features that are likely to be synapomorphies - almost whatever the immediate sister taxon to monocots might be - are in bold. However, if Ceratophyllaceae are sister to monocots, synapomorphies like the herbaceous habit, absence of vascular cambium, etc., will move down a node.
ACORALES Reveal
Inflorescence a spadix, ebracteate, with spathe; ovules atropous; endosperm copious, perisperm +. - 1 family, 1 genus, 2-4 species.
ACORACEAE Martinov - Leaves equitant, isobifacial; peduncle with two separate vascular systems; anther thecae with confluent dehiscence, pollen sulcus lacking ectexine. - 1/2-4. E. America to East and South East Asia.
ALISMATALES + PETROSAVIALES + [DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: ethereal oils 0; raphides +, druses 0; leaf ptyxis variants of supervolute-curved; endothecium develops directly from undivided outer secondary parietal cells, endexine 0, carpels plicate, (septal [epithelial] nectaries +); endosperm nuclear/helobial.
ALISMATALES Dumortier
Starch grains pteridophyte-type, amylophilic; A extrorse, tapetum amoeboid, cells uninucleate, carpels with completely unfused canals, styles separate; endosperm helobial; embryo large; seedling with hypocotyl and root well developed. - 14 families, 166 genera, 4490 species.
ARACEAE Jussieu - Inflorescence terminal, densely spicate [spadix], subtended by an inflorescence bract [spathe], bracts 0; pollen endexine spongy. - 106/4025. Mostly tropical.
Gymnostachydoideae + Orontioideae: ovules atropous.
1. Gymnostachydoideae Bogner & Nicolson - Nucellus not fully covered by integument. - 1/1: Gymnostachys anceps. E. Australia.
2. Orontioideae Mayo, Bogner & Boyce - 3/6. Temperate East Asia, W. and E. North America.
3. Lemnoideae Engler - Thallus-like, stemless, floating aquatic herbs; collenchyma and bundle fibers 0; vessels 0; P 0, A 1, monothecal, or 2, pollen ulcerate, spiny, G 1, embryo sac Allium-type, stigma funneliform; fruit an achene [sort of]; seed operculate; endosperm cellular, starchy, embryo undifferentiated. - 5/37. World-wide.
4. Pothoideae Engler - Stem usu. aerial; styloids +; pseudopetiole apically geniculate; crystals often surrounding the embryo. - 16/1260.
4a. Potheae Engler - 4/900. Tropical America, Madagascar to South and Southeast Asia, Malesia and N.E. Australia.
4b. Monstereae Schott - 12/360. Tropical South and Southeast Asia to the Pacific, South America (Africa).
5. Lasioideae Engler - Inflorescence flowering basipetally; pollen sulcus with ectexine lamella and thick bilayered endexine [outer: flakes or lamellae; inner: spongy]. - 10/58. India and Southeast Asia to the Pacific, also Africa (Lasimorpha).
Calloideae + Aroideae: ?
6. Calloideae Endlicher - P 0, pollen dicolpate. - 1/1: Calla palustris. Circumboreal.
7. Aroideae - Plants monoecious (dioecious); spathe differentiated into tube plus blade, pollen extruded in strands. - 70/2670. Tropical and warm temperate.
7a. Zamioculcadeae Engler - Leaves compound, petiole geniculate; staminate flowers: pollen zonasulcate, columellae winding, forming a sort of internal tectum as well as the external tectum, endexine lamellate, intine thin. - 2/6. Africa, Kenya to Natal.
7b. The rest - Laticifers +; P 0, A connate, connectives thick, pollen inaperturate, ektexine thin, often lacking sporopollenin, endexine thick, spongy, intine massive. - 68/2665. Tropical and warm temperate.
Tofieldiaceae + [Hydrocharitaceae + Scheuchzeriaceae groups]: carpels free.
TOFIELDIACEAE Takhtajan - Stomata anomocytic; leaves equitant, isobifacial; flowers subtended by a "calyculus"; A introrse to latrorse, tapetum secretory, microsporogenesis simultaneous [tetrads tetrahedral], pollen disulcate, nectaries +, ovules unitegmic; seeds with appendage(s). - 3-5/27. S.E. U.S.A., N.W. South America, N. temperate.
Hydrocharitaceae + Scheuchzeriaceae groups: plant aquatic (with floating stems); stem with lacunae; raphides and druses 0; hairs 0; bulliform cells 0 [?this level]; pollen grains trinucleate; endosperm 0; seedling collar and collar rhizoids +.
[Hydrocharitaceae + Butomaceae] [Alismataceae + Limnocharitaceae]: apical meristems of vegetative axes bifurcating; C-glycosyl flavones +; inflorescence scapose, determinate, bracteate; P = K + C, both whorls with many traces, (stamen pairs +), carpels plicate; seeds exotestal.
Hydrocharitaceae + Butomaceae: ?
HYDROCHARITACEAE Jussieu - G inferior; fruit fleshy, dehiscent; endotegmen with tuberculate inner wall alone persisting. - 18/116. World-wide.
1. Hydrocharitoideae Eaton - Leaves broad, vascular bundles inverted, pseudopetiole +, ligules basal; exotestal cells much enlarged. - 2/5. Temperate and subtropical.
Stratiotoideae +{ Anacharidoideae + Hydrilloideae: roots unbranched.
2. Stratiotoideae Luersson - Leaves spiny, in rosettes. - 1/1. Eurasian.
Anacharidoideae + Hydrilloideae: leaves submerged.
3. Anacharidoideae Thomé - 7/38. Tropical to temperate, esp. America.
4. Hydrilloideae Luersson - 8/61. Tropical and subtropical, especially Old World; Naias subcosmopolitan.
BUTOMACEAE Mirbel - Leaves three-angled. - 1/1. Temperate Eurasia.
Alismataceae + Limnocharitaceae: plant with latex; stomata with parallel divisions; leaves involute, with pseudopetiole, midrib, cross veins and an apical subepidermal pore; C thin, more or less crumpled in bud, evanescent, A centrifugal, pollen pantoporate, spinose, G free or connate basally, ovules tenuinucellate, embryo sac tetrasporic, nucellar cap +; fruit a follicle; embryo strongly curved, white; x = 7, 8.
ALISMATACEAE Ventenat - 12/81. Pantropical, also temperate.
LIMNOCHARITACEAE Cronquist - 3/7. Pantropical.
Scheuchzeriaceae + Aponogetonaceae + Juncaginaceae + Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: primary root poorly developed; P members with a single trace.
SCHEUCHZERIACEAE F. Rudolphi - 1/1. N. Temperate to Arctic.
APONOGETONACEAE J. Agardh - 1/43. Old World, esp. South Africa, largely tropical and warm temperate.
Juncaginaceae + Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: sulphated phenolic acids +, leucanthocyanins, flavones 0; leaf ± linear, base auriculate, ligulate; P 0 or reduced [as "retinaculum", "abaxial outgrowth" of A], anthers sessile, pollen inaperturate, nectary 0, carpels with complete postgenital fusion [sampling!].
JUNCAGINACEAE Richard - 4/15. Cosmopolitan.
Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: aquatics; rhizome with endodermis; epidermis chlorophyllous; water pollination; carpels ascidiate [sampling!], 1 apical atropous ovule/carpel; fruit ± drupaceous; embryo with massive elongated hypocotyl, also prominent in seedling, collar or base of hypocotyl much enlarged.
Posidoniaceae + Ruppiaceae + Cymodoceaceae: ± marine; vessels 0; stomata 0; leaves 2-ranked; pollen ± filiform.
POSIDONIACEAE Hutchinson - Plant monopodial, with copious unlignified fiber strands; inflorescence pedunculate, branched-racemose; G 1, stigma sessile, ornate; pericarp fleshy. - 1/9. Mediterranean, temperate Australia.
Ruppiaceae + Cymodoceaceae: leaves serrulate; A 2.
RUPPIACEAE Horaninow - Pollen grains arcuate, stigma sessile, capitate; stone with operculum. - 1/1-10. More or less world-wide.
CYMODOCEACEAE N. Taylor - Anther with filament, pollen grains much elongated. - 5/16. More or less tropical (to warm temperate), Australia in particular.
Zosteraceae + Potamogetonaceae: leaf with apical pore, (sheath closed); plant mono- or dioecious; P and A pair with single vascular trace.
ZOSTERACEAE Dumortier - Inflorescence with spathe and spadix, spadix axis flattened, flowers two ranked, alternating on adaxial surface, bracts 0; A 1, with filament, thecae separate, deciduous, joined by divided connective, pollen filiform, G [2]. - 2/14. Temperate to subtropical.
POTAMOGETONACEAE Reichenbach - 7/102. Worldwide, esp. temperate.
PETROSAVIALES + [DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: Starch grains simple, amylophobic; stomata anomocytic; colleters 0; endosperm nuclear.
PETROSAVIALES Takhtajan
Stem with a ring of bundles; sieve tube plastids also with polygonal protein crystalloids; microsporogenesis simultaneous, septal nectaries +; fruit a follicle.
PETROSAVIACEAE Hutchinson* - T whorls slightly differentiated [outer somewhat smaller], integumentary obturator +. - 2/4. China, Japan, W. Malesia. 1 family, 2 genera, 4 species.
[DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: Inflorescence type?
[DIOSCOREALES + PANDANALES]: ?
DIOSCOREALES J. D. Hooker
Vascular bundles in rings; leaves spiral; flowers or inflorescence with glandular hairs; styles initially at most weakly connate, short; T persistent in fruit; endotegmen tanniniferous; embryo short. - 5 families, 21 genera, 1037 species.
NARTHECIACEAE Bjurzon - Raphides 0, druses +; ovary ± inferior in flower; endosperm helobial. - 4-5/41. N. temperate, Venezuela and Guiana.
Taccaceae + Thismiaceae + Burmanniaceae + Dioscoreaceae: stem with endodermis; T tube broad, ovary inferior, stigma lobes ± bilobed; fruit winged; exo- and endotesta tanniniferous.
Taccaceae + Thismiaceae: A reflexed in flower, septal nectaries 0, placentation parietal.
TACCACEAE Berchtold & J. Presl Hairs with multicellular stalk row, a head, and then another cell row; petiole bundles in ring; leaves basal, petiolate; inflorescence scapose, umbellate, of groups of cinicinni, bracteate, long filiform bracts among the flowers. - 1/12. Pantropical, esp. Malesian-Pacific.
THISMIACEAE Agardh - Plants ± myco-heterotrophic; stomata 0; leaves reduced to scales; T whorls often differentiated, both well developed, A with intrastaminal lobes, connate postgenitally into a ring, pollen porate, ovules tenuinucellate, style connate early in ontogeny; fruit irregularly dehiscent, P usually circumscissilely dehiscent. - 4/31. Widely scattered, (sub)tropical.
Burmanniaceae + Dioscoreaceae: ?
BURMANNIACEAE Blume - Raphides 0; outer T larger, enclosing the inner, A 3, opposite inner T, thecae separated by connective, tranversely dehiscent, ovules tenuinucellate, stigma infundibular. - 9/95. Largely tropical, esp. American.
DIOSCOREACEAE R. Brown - Vessels in cauline bundles interrupted at nodes by tracheids, sieve tubes similarly interrupted; leaves petiolate, with midrib and reticulate fine venation, petiole pulvinate at both ends; fruits winged, seeds winged; endotestal cells thick-walled, with crystals, exotegmen thickened. - 3/805. Largely tropical.
PANDANALES Lindley
Nucellar cap +; embryo minute, endosperm type? - 5 families, 36 genera, 1345 species.
VELLOZIACEAE Hooker - Xeromorphic; sieve tube plastids in the stem <1 µm across; P large, ovules tenuinucellate, style long. - 9/240. South America and Africa-Madagascar to Arabia, South East Asia.
Acanthochlamys - Basal part of scape with a root-like vascular cylinder; leaf with large sheathing basal ligule; midrib with back-to-back vascular bundles; inflorescence compound capitate, scapose; A bisporangiate/unithecal, septal nectaries 0. - 1/1. S.E. Tibet and S.W. China.
Vellozia, etc. - Sieve tube plastids with angular crystals and other loosely-packed crystals; transfusion tracheids in leaf bundles; anthers long; inflorescence more or less sessile. - 8/240. South America and Africa-Madagascar (to Arabia).
Triuridaceae + Stemonaceae [Pandanaceae + Cyclanthaceae]: septal nectaries 0.
TRIURIDACEAE Gardner - Echlorophyllous myco-heterotrophic herbs; vessels 0; stem with a ring of bundles; stomata 0; inflorescence a raceme; T valvate, carpels free, 1 (2) basal tenuinucellate ovule/carpel. - 8/48. Pantropical.
Stemonaceae + Pandanaceae + Cyclanthaceae: flowers other than 3-merous; placentation parietal, style 0.
STEMONACEAE Caruel - Leaves with petiole and well-developed fine cross veins; anther connective expanded; testa multilayered, seeds with longitudinal ridges. - 4/27. China and Japan to Australia, S.E. U. S. A.
Pandanaceae + Cyclanthaceae: stem vascular bundles compound; stomata tetracytic; inflorescence bracts conspicuous, flowers congested, sessile, imperfect; staminate flowers: stamens usu. several-many, pollen porate; carpellate flowers: fruit an indehiscent syncarp; endotesta well developed; cotyledon not photosynthetic, all seedling internodes ± elongated.
PANDANACEAE R. Brown - Woody, not rhizomatous; sieve tube plastids also with peripheral fibers; leaves M-shaped when mature, spiny; inflorescence bracts usually colored. - 3/885. W. Africa to the Pacific.
CYCLANTHACEAE A. Richard - Leaves petiolate, basically simple, plicate or variants, often divided deeply; ovules tenuinucellate, micropyle bistomal. - 12/225: Central and tropical South America.
Cyclanthoideae - Subepidermal sclereids +; non-articulated laticifers +; lysigenous air spaces with transverse septae +; inflorescence with whorls of staminate and carpellate "flowers", P 0; staminate flowers: A in 4 rows per whorl, connate basally; carpellate flowers: ovary cavity with closely-set placentae; fruit dry, splitting down the middle of the carpellary annulus; seeds embedded in mucilage. - 1/1. Central and N. South America, the Lesser Antilles.
Carludovicioideae - T 10-30, with abaxial gland; inflorescence with spathe, flowers in spirals; staminate flowers: T in one or two whorls, 6+, filaments swollen basally; carpellate flowers: 4(-8)-merous, staminodes opposite P, long-filiform; fruit baccate. - 11/225. Central and tropical South America.
LILIALES + ASPARAGALES + COMMELINIDS: ?
LILIALES Perleb
Plants geophytes; leaves elliptical, fine venation reticulate, base not sheathing; T large, (spotted), A extrorse, tepal nectaries alone, many tenuinucellate ovules/carpel, nucellar cap +; tegmen with cellular structure; endosperm with thick pitted walls, hemicellulosic; mitochondrial sdh3 gene lost. - 11 families, 67 genera, 1558 species.
CORSIACEAE Beccari - Echlorophyllous myco-heterotrophic herbs; leaf sheath closed; flowers single, monosymmetric, held inverted, median T of outer whorl standard-like ["labellum"], pollen porate, G inferior, funicle long; embryo undifferentiated. - 3/30. S. China, S. South America, Papuasia-Australia.
CAMPYNEMATACEAE Dumortier - Fibrous leaf bases persistent; tepals not spotted, ovules crassinucellate; T enlarging, persistent in fruit. - 2/4. New Caledonia and Tasmania.
<[Petermanniaceae [Colchicaceae [Alstroemeriaceae + Luzuriagaceae]]] Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]: (fruit a berry).
Petermanniaceae [Colchicaceae [Alstroemeriaceae + Luzuriagaceae]]: primary root of seedling well developed.
PETERMANNIACEAE Hutchinson - Rhizome woody; stem with prickles and leaf-opposed tendrils; leaves subpetiolate, with midrib and parallel secondary veins; tepals not spotted, tapetum amoeboid, G inferior, placentation parietal; fruit a berry. - 1/1. Central part of the E coast of Australia.
Colchicaceae [Alstroemeriaceae + Luzuriagaceae]: ?
COLCHICACEAE de Candolle - 15/245. Temperate to tropical, but not in South America.
1. Burchardieae (Bentham) J. C. Manning & Vinnersten - Rhizome short, vertical, with papery scales; leaves sheathing; capsule septicidal. - 1/?. Australia.
Uvularieae [Tripladenieae [Colchiceae [Iphigenieae + Anguillarieae]]]: flowers axillary.
2. Uvularieae Meisner - Rhizomes +. - 2/15. W. and E. North America, East Asia to W. Malesia.
Tripladenieae [Colchiceae [Iphigenieae + Anguillarieae]]:
3. Tripladenieae Vinnersten & J. C. Manning - Rhizome +. - 3/5. Australia and New Guinea.
Colchiceae [Iphigenieae + Anguillarieae]: tunicated corm +; alkaloids with a 7-C tropolone ring +; leaves sheathing.
4. Colchiceae Reichenbach - Nectaries median on tepal or on stamen base. - 5/170. Africa, Europe, Central to tropical South East Asia.
Iphigenieae + Anguillarieae: ?
5. Iphigenieae Hutchinson - Nectaries 0. - 2/10. Old Wold Tropics, South Africa..
6. Anguillarieae D. Don - Inflorescence racemose or spicate, bracts 0. - 2/38. Africa, Australia.
Alstroemeriaceae + Luzuriagaceae: leaves resupinate; testa and tegmen thin-walled; karyotype bimodal.
ALSTROEMERIACEAE Dumortier - Leaves ± resupinate; inflorescence terminal, a condensed thyrse, bracteoles lateral; flower monosymmetric, median member of the outer T whorl adaxial, inner whorl often with nectariferous claw, A latrorse, centrifixed, G inferior. - 3/165. Tropical and temperate Central and South America.
LUZURIAGACEAE Lotsy - Stems perennial; leaves 2-ranked, resupinate; tepals not spotted; fruit a berry. - 2/5. Peru to Tierra del Fuego, Falkland Islands, New Zealand and Australia (New South Wales to Tasmania).
MELANTHIACEAE Batsch - 16/170. N. temperate, esp. East Asia and E. North America.
1. Meliantheae - Bulbous; highly oxygenated esterified C-nor-D homosteroidal alkaloids; anthers kidney bean-shaped, opening by valves; capsule septicidal [ventricidal]; n = 8. - 7/100. North Temperate, Schoenocaulon to Peru.
Helionadeae + Chionographideae: Calcium oxalate crystals cuboidal; bracts 0; pollen intectate.
2. Helionadeae - Pollen spinulate; seeds linear, long-caudate at both ends; n = 17. - 1/9. E. North America, East Asia.
3. Chionographideae - Flowers often imperfect; T with 1 nerve, pollen 4-porate, with clavate processes; capsule septicidal; seeds winged (at one end); n = 12. - 2/6. E. North America, East Asia.
Xerophylleae + Parideae: anther thecae distinct.
4. Xerophylleae - Plant ± bulbous; leaf long-linear, xeromorphic; petal nectaries 0; n = 15. - 1/2. North America.
5. Parideae - Leaves whorled, with (petiole), midrib, and broad blade, venation reticulate; flowers single, terminal; P = K + C, embryo sac bisporic; embryo minute, undifferentiated; n = 5, chromosomes heteromorphic, 6-40+µm long. - 4/ca. 80. North Temperate.
[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]: leaves broad, with reticulate venation; fruit a berry.
Philesiaceae + Rhipogonaceae: stem fructans 0; cuticular wax with parallel platelets; leaves with petiole, lamina, midrib; tepals not spotted, ovules crassinucellate; testa disintegrates.
PHILESIACEAE Dumortier - Pollen inaperturate, spinose, placentation intrusive parietal. - 2/2. Chile.
RHIPOGONACEAE Conran & Clifford - Stem prickly; leaves opposite; flowers small, tapetum amoeboid, 2 ovules/carpel. - 1/6. New Zealand to New Guinea.
<Smilacaceae + Liliaceae: ?
SMILACACEAE Ventenat - Stem prickly; leaves with lateral ligules and/or thorns or tendrils, with petiole, lamina, midrib; inflorescence umbellate; flowers small, tepals not spotted, pollen inaperturate, spinulose, 1 ovule/carpel. - 2/315. Pantropical to temperate.
LILIACEAE Jussieu - Bracteoles lateral; T often spotted, anthers often centrifixed. - 16/635: North Temperate, especially East Asia and North America.
Lilioideae - Embryo sac tetrasporic; capsule septicidal. - 11/535. (Cold) temperate North America, East Asia.
Calochortoideae - 5/100. N. (cool) temperate, esp. East Asia and E. and W. North America.
ASPARAGALES + COMMELINIDS: ?
ASPARAGALES Bromhead
Chelidonic acid; anthers longer than wide, tapetal cells bi- to tetranuclear, microsporogenesis simultaneous, style single; seeds exotestal, tegmen not persistent; endosperm helobial; mitochondrial sdh3 gene lost. - 16-24 families, 1122 genera, 26071 species.
ORCHIDACEAE Jussieu - Mycorrhizal herbs, protocorms myco-heterotrophic; SiO2 bodies [stegmata] in bundle sheaths; inflorescence racemose; flowers monosymmetric, resupinate, labellum + [median petal of inner whorl], A 3 [median of outer whorl and laterals of inner whorl], basally adnate to style, tapetal cells uninucleate, septal nectaries 0, G inferior, placentae branched, very many tenuinucellate ovules/carpel, funicle not vascularised; seeds minute; endosperm barely developing, embryo minute, undifferentiated. - 788/18000-20000. World-wide.
1. Apostasioideae - 2/16. Sri Lanka, N.E. India to N.E. Australia, Japan.
Vanilloideae + Cypripedioideae + Orchidoideae + Epidendroideae: C-glycosyl flavones, (saponins), 6-hydroxy flavonols +; leaves spiral or 2-ranked; flowers strongly monosymmetric; labellum strongly differentiated, the style and A almost completely congenitally fused [gynostemium], anthers to 2x as broad as long, pollen sticky, placentation parietal, stigma asymmetrical; ovules not fully developed at pollination, fertilisation may take some months; seeds dispersed before maturity of embryo; radicle 0.
2. Vanilloideae Szlachetko - A 1 [median member of outer whorl], staminodes 2 [from inner whorl]. - 15/230. Pantropical, esp. Asia; Australia, some N. America.
Cypripedioideae + Orchidoideae + Epidendroideae: ?
3. Cypripedioideae - 2 abaxial T of outer whorl connate, labellum saccate, A 2 [from inner whorl], staminode 1 [median member of outer whorl]. - 5/130. Mostly (warm) temperate N. hemisphere, East Malesia and tropical South America (S. India).
Orchidoideae + Epidendroideae: floral primordium tranversely elliptic-oval; labellum initiated first, A 1 [median member of outer whorl], pollen forming pollinia that are variously attached to sticky viscidium, pollinarium stalk variously formed from part of the anther [caudicula], epidermis of the rostellum [tegula] or apex of the rostellum [hamulus], microsporogenesis simultaneous, pollen usu. in tetrads, porate or ulcerate, median carpel developed before the others, rostellum [ridge, part of median stigmatic lobe, viscidium is also part of it] +; T not abscising; tegmen not persisting.
4. Orchidoideae Lindley - Leaves soft, deciduous. - 208/3630. World-wide, esp. temperate.
5. Epidendroideae - Anthers incumbent. - 650/18000. Especially tropical, some temperate, rather poorly developed in Australia.
Boryaceae et al. + Ixoliriaceae, etc. + Doryanthaceae + Iridaceae + Xeronemaceae + Hemerocallidaceae, etc., + Alliaceae, etc., + Asparagaceae, etc.: stem fructans +; cuticle wax crystals as parallel platelets; (T ± connate, A inserted on T tube); seeds exotestal, with phytomelan.
Boryaceae + Blandfordiaceae + Lanariaceae + Asteliaceae + Hypoxidaceae: ovules with hypostase, embryo sac with chalazal constriction.
BORYACEAE M. W. Chase, Rudall & Conran - 2/12. Australia.
Blandfordiaceae + Lanariaceae + Asteliaceae + Hypoxidaceae: ovules with parietal cell and nucellar cap.
BLANFORDIACEAE R. Dahlgren & Clifford - Pollen trichotomosulcate, G stipitate, septal nectaries external. - 1/4. E. Australia.
Lanariaceae + Asteliaceae + Hypoxidaceae: hairs multicellular, often branched; stomata paracytic; lamina with distinct midrib; micropyle bitegmic.
LANARIACEAE R. Dahlgren & A. E. van Wyk - 1/1. Cape Province, South Africa.
Asteliaceae + Hypoxidaceae: rosette-forming or caespitose; flavonols +; mucilage canals +; endosperm thin-walled; cotyledon not photosynthetic, ligule long.
ASTELIACEAE Dumortier - Ovary with mucilage. - 2-4/36. New Zealand to New Guinea, Pacific Islands E. to Hawaii, Chile, the Mascarenes.
HYPOXIDACEAE R. Brown - Leaf bases persisting; inflorescence axis compressed; septal nectary 0, tapetum plasmodial; seed with prominent raphe. - 7-9/100-220. Seasonal tropics, esp. southern Africa (temperate).
Ixioliriaceae + Tecophilaeaceae: cormose; leaves spiral; outer T mucronate to aristate, T tube short.
IXOLIRIACEAE Nakai - 1/3. Egypt to Central Asia.
TECOPHILAEACEAE Leybold - Anthers ± poricidal, pollen operculate. - 8/23. Africa, Chile and U.S.A (California).
DORYANTHACEAE R. Dahlgren & Clifford - Huge sub-bulbous tufted perennial; styloids +, raphides 0; older leaves with dry threads at apex; A centrifixed, connective massive, endothecium thick, pollen trichotomosulcate, G inferior. - 1/2. E. Australia.
Iridaceae + Xeronemataceae + Hemerocallidaceae, etc., + Alliaceae, etc., + Asparagaceae, etc.: ?
IRIDACEAE Jussieu - Styloids +; leaves equitant, isobifacial; inflorescence a riphidium; T ± free, A 3, extrose, stigmas at the edges of the complex/expanded style. - 67/1800. World-wide.
1. Isophysidoideae Takhtajan - 1/1. Tasmania.
Iridoideae [Nivenioideae + Crocoideae]: xanthone [mangiferin] +; G inferior; endosperm usually nuclear.
2. Iridoideae - T nectaries, oil glands or oil hairs + [for most of the clade]. - 30/765. Worldwide, but esp. the spine of Central and South America.
Nivenioideae + Crocoideae: extra codon in rps4 gene.
3. Nivenioideae Goldblatt - Inflorescence biseriate, of two terminal monochasia subtended by inflorescence bracts; flowers blue, T connate; seeds shield-shaped, testa 2-3 cells thick, exotesta transparent, unthickened, (exfoliating), exotegmen well developed. - 5/65. Africa, especially the Cape, and Madagascar.
4. Crocoideae G. T. Burnett - Plant with corms: leaves with pseudomidrib (not Pillansia), sheath closed; inflorescence spicate; flowers obliquely monosymmetrical (polysymmetrical), T connate or not, pollen exine tectate, perforate-scabrate, aperture operculate, ovules campylotropous, chalazal hypostase prominent. - 28/995. Overwhelmingly southern African, to Europe, Madagascar and Central Asia.
Xeronemataceae [Hemerocallidaceae, etc. [Alliaceae, etc., + Asparagaceae, etc.]]: mitochondrial rpl2 gene lost.
XERONEMATACEAE M. W. Chase, Rudall & Fay - 1/2. New Zealand (Poor Knights Island) and New Caledonia.
Hemerocallidaceae, etc., + Alliaceae, etc., + Asparagaceae, etc.: (pedicels articulated); septal nectaries infralocular, ovary superior, microsporogenesis successive.
Hemerocallidaceae + Xanthorrhoeaceae + Asphodelaceae [= Xanthorrhoeaceae s.l.]: anthraquinones +; styloids +; cotyledon not photosynthetic.
HEMEROCALLIDACEAE R. Brown - Leaves with unifacial keel. - 19/85. Papuasia to New Zealand and the Pacific, esp. Australia, also Europe to Asia, Malesia, India, Madagascar, and Africa.
Xanthorrhoeaceae + Asphodelaceae: (secondary thickening +;) A not adnate to T, hypostase +; seeds angled.
XANTHORRHOEACEAE Dumortier - Stem thick, woody; plant resiniferous; raphides 0; layer of sclerenchyma below epidermis; stomata paracytic; leaves unifacial, not sheathing; inflorescence spicate, of congested cymes; T = 3 dry + 3 subpetaloid, pollen extended sulcate; embryo transverse to long axis of seed. - 1/30. Australia.
ASPHODELACEAE Jussieu - Parenchymatous cells in the inner bundle sheath adjacent to the phloem. - Ca 15/785. Africa, esp. South Africa, also New Zealand (Bulbinella) and the Mediterranean to Central Asia.
Alliaceae, etc., + Asparagaceae, etc.: microsporogenesis successive [possible place].
Agapanthaceae [Alliaceae + Amaryllidaceae]: flavonols, saponins +; laticifers +; mucilage cells +; leaves 2-ranked; inflorescence scapose, umbellate, with scarious spathe, inflorescence bracts 2 (or more - external).
AGAPANTHACEAE F. Voigt - 1/9. South Africa.
Alliaceae + Amaryllidaceae: sympodial tunicate-bulbous geophytes with contractile roots.
ALLIACEAE Borkhausen - Cysteine-derived sulphur compounds +; ovules tenuinucellate. - 13/795. Mainly South America, but Allium esp. N. Temperate Eurasia.
1. Allioideae Herbert - Bulbs lacking starch; A often with winged filaments, style solid, gynobasic; endosperm cellular, embryo curved. - 2/260-690. North temperate, often seasonally dry, especially Central Asia, scattered in Africa.
Tulbaghioideae + Gilliesioideae: bulbs with starch; endosperm helobial.
2. Tulbaghioideae (Meisner) M. F. Fay & M. W. Chase - Corona massive, A sessile. - 1/22. Southern Africa.
3. Gilliesioideae (Lindley) Arnott - Corona +/0; embryo short. - 10/80. South America.
AMARYLLIDACEAE J. Saint-Hilaire - Distinctive alkaloids +; G inferior. - 59/800+. Tropical (temperate), esp. South America and Africa, also Mediterranean.
1. Amaryllideae J. Saint-Hilaire - Extensible [helically-thickened] fibers in leaf; pollen bisulcate, exine gemmate, with scattered spinules, intectate-columellate, ovules unitegmic, embryo sac Allium type; seeds water-rich, non-dormant, testa to 25 cells thick, chlorophyllous, with stomata, or ± collapsed, with a corky layer, endosperm chlorophyllous, phytomelan 0; embryo chlorophyllous. - 11/146. SubSaharan, especially South Africa, Crinum Pantropical.
Cyrtantheae, etc.: bundle sheath cells parenchymatous.
2. Cyrtantheae Salisbury - Scape lacking sclerenchymatous ring, subepidermal collecnchyma +; 1-layered rhizodermis +, velamen 0. - 1/50. Africa, especially the south.
3. Calostemmateae D. & U. Müller-Doblies - 2-3 ovules/carpel; embryo germinates precociously; fruit indehiscent; phytomelan 0; n = 10. - 2/4. Australia, Malesia.
4. Haemantheae (Pax) Hutchinson - 1-layered rhizodermis +, velamen 0; scape lacking sclerenchymatous ring, subepidermal collenchyma +. - 6/80. Tropical Africa, mostly in the South.
Eurasian Clade: seeds subglobose, turgid.
5. Lycoridae Traub - 2/26. Temperate to subtropical East Asia to Iran.
6. Galantheae Salisbury - Elaiosome +. - 8/31. Europe to N. Africa, the Crimea and the Caucasus.
7. Pancratieae Salisbury - Staminal tube toothed; n = 11. - 1/20. Mediterranean, southern Asia, to sub-Saharan Africa./p>
8. Narcisseae Endlicher - Inflorescence bracts basally connate; elaiosome +. - 2/58: Narcissus (50). Europe to W. Asia and N. Africa.
Andean + extra-Andean clade: 1-layered rhizodermis +, velamen 0; scape lacking sclerenchymatous ring, subepidermal collenchyma +; bracts obvolute; (seeds flat, horizontally stacked).
9. Hippeastreae (Pax & Hoffmann) Huchinson - Inflorescence bracts often connate basally (along one side); flowers monosymmetric, A declinate, of varying lengths; seeds flattened. - 11/218. S.E./S.W. U.S.A., the Caribbean, and Central and South America.
Eustephieae + Hymenocallideae + Stenomesseae + Eucharideae: no palisade leaf mesophyll; x = 23 (tetraploid).
10. Eustephieae (Pax) Hutchinson - A of two lengths; seeds flattened, winged. - 3/15: C. Andes.
Hymenocallideae + Stenomesseae + Eucharideae: ?
11. Hymenocallideae (D. & U. Müller-Doblies) Meerow - Pollen grains auriculate [the two ends narrowed, with different sculpture]; testa thick, spongy, vascularised, phytomelan 0/ - 3/65. S.E. U.S.A., the Antilles, Central America to Bolivia.
12. Stenomesseae Traub - Leaves petiolate, lorate; staminal cup + (0); seeds flattened, obliquely winged. - 8/62. Andean South America S. to Bolivia, Costa Rica.
13. Eucharidae (Pax) Hutchinson - Leaves petiolate; seeds globose, turgid, coat lustrous. - 4/28. Central America, the Andes S. to Bolivia.
[Aphyllanthaceae [Themidaceae + Hyacinthaceae] Agavaceae] [Laxmanniaceae [Asparagaceae + Ruscaceae]] [= Asparagaceae s.l.]: inflorescence racemose.
Aphyllanthaceae [Themidaceae + Hyacinthaceae] Agavaceae: ?
APHYLLANTHACEAE Burnett - Stems photoysnthetic, with stomata in bands down its length; leaves as non-photosynthetic scales; inflorescence scapose, flowers multibracteolate; T marcescent, pollen spiraperturate, 1 ovule/carpel. - 1/1. W. Mediterranean.
Themidaceae + Hyacinthaceae: leaves spiral; pedicels bracteate; cotyledon not photosynthetic.
THEMIDACEAE Salisbury - Inflorescence scapose. - 12/62. S.W. North America, to British Columbia and Guatemala.
HYACINTHACEAE Borkhausen - Plant bulbous geophytes, roots often contractile; polyhydroxyalkaloids, flavones, flavone C-glycosides +; little sclerenchyma in the leaf; mucilage cells +; inflorescence scapose. - 41-70/770-1000. Predominantly Old World, esp. S. Africa, and the Mediterranean to Central Asia and Burma.
1. Oziroëoideae Speta - A basally connate and adnate to C. - 1/5. Western South America.
Ornithogaloideae + Urgineoideae + Hyacinthoideae: rhexigenetic lacunae +; also styloids +.
2. Ornithogaloideae Speta - Cardenolides +; protein crystals in nucleus. - 1/200. Europe, W. Asia, Africa.
3. Urgineoideae Speta - Bufadienolids +; bracts spurred [as small leaves in Bowiea]; testa brittle, not tightly adherent to endosperm. - 2(-3?)/105. Mainly Africa, Madagascar, the Mediterranean to India.
4. Hyacinthoideae Link - Homoisoflavanones +; seeds often rounded.
4a. Pseudoprospereae Manning & Goldblatt - 1/1. E. South Africa.
4b. Massonieae Baker - Ca 10/235. Africa S. of the Sahara, Ledebouria to India.
4c. Hyacintheae - 21/265. Europe to the Mid (Far) East, North Africa.
AGAVACEAE Dumortier - 23/637. More or less world-wide, esp. S.W. North America, few in Malesia, N. Australia, not cold temperate or New Zealand, etc.
Laxmanniaceae [Asparagaceae + Ruscaceae]: endosperm helobial, thick-walled, pitted, hemicellulosic.
LAXMANNIACEAE Bubani - 14-15/178. Predominantly Australian, also Madagascar, India and South East Asia to the Pacific, and South America.
1. Lomandra group - Lamina with sclerenchymatous ribs extending from the inner sheath of the vascular bundle to the surface, outer bundle sheath with enlarged cells; leaves 2-ranked, flat or curved; 1-2 ovules/carpel, nucellus with axially oriented central conducting passage; phytomelan 0, testa thin, tegmen brown, collapsed, cellular. - 5/65. Australia, New Guinea, New Caledonia.
2. Laxmannia group - Plant with (ecto) vesicular-arbuscular mycorrhizae; storage roots +; leaves spiral; nucellus with axially oriented central conducting passage; endosperm cell walls thin. - 8/92. - South East Asia to Australia, New Zealand and the Pacific.
3. Cordyline group - Rosette herbs to trees; storage roots +; stomata paracytic; leaves spiral; endosperm cell walls thin. - 2/17. India to the Pacific and New Zealand, tropical America.
Asparagaceae + Ruscaceae: flowers small?
ASPARAGACEAE Jussieu - Embryo sac curved, nucellar epidermal cells enlarged. - 2/165-295. Old World, but not the Antipodes (except N. Australia), Mexico.
RUSCACEAE Sprengel - 26/475. N. hemisphere, esp. Southeast Asia-Malesia (Convallariaceae s. str.), Europe and the Near East (Ruscaceae s. str.), S.W. North America (Nolinaceae s. str.), Africa, esp. the Cape and S.W. (Eriospermaceae s. str.).
COMMELINIDS
Unlignified cells walls with UV-fluorescent ferulic and coumaric acids; SiO2 bodies in leaves; stomata para- or tetracytic, (cuticular waxes as aggregated rodlets [looking like a scallop of butter]); inflorescence bracteate; (P fully bicyclic) [A adnate to C/inner P], pollen starchy; embryo short, broad.
DASYPOGONACEAE Dumortier - 2 peripheral phloem strands in foliar bundles; 1 ascending ovule/carpel. - 4/16. Southern Australia.
Dasypogon + Calectasia - Hairs branched; fruit indehiscent; tegmen collapsing, massive storage nucellus with starch [below embryo sac]. - 2/14. S.W. Australia, Victoria.
Kingia + Baxteria - Rhizome 0; raphides 0; inflorescence surrounded by bracts, stalk bracteate; pollen extended sulcate-unipantocolpate. - 2/2. S.W. Australia.
ARECALES Bromhead
Plant woody, monopodial, not rhizomatous; cuticular waxes as aggregated rodlets, stomata tetracytic; leaves spiral, petiolate, reduplicate-plicate, pinnately pseudocompound; flowers ± sessile, 1 apotropous ovule/carpel. - 1 family, 189 genera, 2361 species.
ARECACEAE Schultz-Schultzenstein//Palmae Jussieu- SiO2 bodies hat-shaped; leaf with closed sheath; fruit indehiscent; endosperm with hemicellulose. - 189/2361. Humid tropics and subtropics (warm temperate).
1. Calamoideae Beilschmied - Climbers, internodes elongated; fruits with recurved scales. - 22/. Tropical, but esp. Sri Lanka to West Samoa and Fiji.
2. Nypoideae Griffith - Branching dichotomous, plant with horizontal axes; leaves reduplicate; inflorescence with axes adnate to internode above; carpels separate. - 1/1. Malesia (Bengal to Queensland).
Coryphoideae [Ceroxyloideae + Arecoideae]: microsporogenesis simultaneous.
3. Coryphoideae Burnett - Leaves induplicate. - 44/ . Africa, esp. South East Asia.
4. Ceroxyloideae Drude
- 8/ . Mostly Central and W. South America, also Madagascar, Florida and the Antilles5. Arecoideae Burnett - 112/ . Pantropical.
POALES + COMMELINALES + ZINGIBERALES: Primary cell wall mostly with glucurono-arabinoxylans; stomata with parallel cell divisions; endosperm starchy.
POALES Small
Vessels also in stem and leaf; SiO2 epidermal; raphides 0; endosperm nuclear, mebryo short to minute; mitochondrial sdh3 gene lost. - 17 families, 997 genera, 18325 species.
Sparganiaceae + Typhaceae: Plant rhizomatous; flavonoids +; SiO2 bodies 0; starch grains pteridophyte-type, amylophilic; leaves 2-ranked; plant monoecious; inflorescences complex; flowers very small, P chaffy, A 1-8, tapetum plasmodial, 8 nuclei/cell, pollen monoulcerate, G pseudomonomerous, nectary 0, 1 pendulous apotropous ovule/carpel, micropyle bistomal, stigma rather elongated; endosperm helobial, cell wall formation in small chalazal chamber before that in large micropylar chamber, perisperm thin, embryo long, slender; x = 15; seedling with hypocotyl and collar hairs.
SPARGANIACEAE Hanin - Leaf sheath not distinct; inflorescence as globose heads; carpellate flowers: antipodal cells multiply after fertilisation. - 1/14. Temperate and Arctic, to New Zealand.
TYPHACEAE Jussieu - Inflorescence densely spicate; carpellate flowers: long hairs on pedicels; endosperm also with oil. - 1/8-13. Temperate and tropical regions worldwide.
BROMELIACEAE Jussieu - Silica bodies solitary; hairs water-absorbing; leaves lacking a sheath; P = K + C, stigma conduplicate spiral. - 57/1400. (Sub)tropical America; one species in W. tropical Africa.
1. Brocchinia - Leaves with stellate chlorenchyma; G ± inferior, septal nectaries above the ovules. - 1/21. South America, the Guyana Highlands.
Rest: cap cells of trichomes dead, septal nectaries below the ovules.
2. Lindmania - Plant dioecious; K contorted, stigma simple-erect. - 1-2/43. South America, the Guyana Highlands.
3. Tillandsioideae Burnett - Epiphytes, roots often for attachment only (0); scales radially symmetrical; ovules with chalazal appendage; seeds with apical and/or basal tufts of hair. - 9/1015. Almost the range of the family in America.
4. Hechtia - Plant dioecious. - 1/51. Texas, Mexico, N. Central America.
5. Navioideae Harms - 5/105. Guyana Highlands, N.E. Brasil.
6. Pitcairnioideae Harms - Scales ± divided; seeds tailed. - 6/515. Mexico to Chile, Pitcairnia feliciana W. Africa.
7. Puya - K contorted, C clawed, tightly spiralled after anthesis; seeds winged. - 1/195. Mountains, etc., Costa Rica and Guyana to Chile and Argentina.
8. Bromelioideae Burnett - Epiphytes, roots often for attachment only; scales irregular peltate; G inferior; fruit baccate. - 31/722. Mexico and the West Indies to Chile.
Rapateaceae + Cyperaceae group + Poaceae group: ?
RAPATEACEAE Dumortier - Mucilage cells +; stomatal guard cells dumbbell-shaped; uniseriate [slime-secreting] colleters +; inflorescence capitate, scapose; P = K + C, anthers with pores, microsporogenesis simultaneous. - 16/94. Tropical South America, in Africa, only Maschalocephalus.
1. Rapateoideae Maguire - Involucral bracts long; 1 ovule/carpel; seeds ovoid-oblongoid. - 3/29. The Guianas to Bolivia and the Matto Grosso.
2. Monotremoideae Givnish & P. E. Berry - 1 ovule/carpel; seeds ovoid-oblongoid, white-granulate [muriculate], with flattened apical appendage. - 4/8. Guiana, upper Rio Negro in Colombia and Venezuela, Maschalocephalus dinklagei in Sierra Leone and Liberia.
3. Saxofridericioideae Maguire - Seeds prismatic, pyramidal, lenticular or crescent-shaped. - 9/54. N. South America, Panama.
Cyperaceae group + Poaceae group: septal nectary 0, style ± strongly divided; mitochondrial sdh4 gene lost.
Thurniaceae + Juncaceae + Cyperaceae: mycorrhizae 0; 3-desoxyanthocyanins [?Cyperaceae], luteolin 5-methyl ether +; trichoblasts from distal cell of pair; air canals in leaves [?= septate aerenchyma]; stem angled, leaves 3-ranked, sheaths closed; flowers small, T scarious, undifferentiated, microsporogenesis simultaneous, pollen in tetrads, porate; seeds testal-tegmic; chromosomes with diffuse centromeres; seedling collar inconspicuous, with rhizoids; deletions in ORF 2280 region.
THURNIACEAE Engler - 2/4. South Africa and Guyana region, Amazonia.
Juncaceae + Cyperaceae: luteolin +; micropyle endostomal; chloroplast rpl23 gene absent.
JUNCACEAE Jussieu - Stems round. - 7/430. Worldwide, esp. Andes (3 endemic genera), S. South America-New Zealand (2 endemic genera).
CYPERACEAE Jussieu - SiO2 bodies conical, attached to walls; 1 basal ovule/flower; 3 bp 5.8S nrDNA insertion, rps14 gene to nucleus, pseudogene remaining in mitochondrion. - 98/4350. World-wide.
1. Mapanioideae - Flowers pseudanthia, with stamens in axils of bracts surrounding carpellate flowers[???]. - 6/140. Largely tropical.
2. Cyperoideae - T = scales, bristles, 0, pollen obovoid, also with 3-6 lateral pores/colpi (pantoporate). - 92/4210. World-wide.
[Mayacaceae [Eriocaulaceae + Xyridaceae]] [[Anarthriaceae [Restionaceae + Centrolepidaceae]] [Flagellariaceae [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]: ovules tenuinucellate; embryo minute.
Mayacaceae + Eriocaulaceae + Xyridaceae: flavonoids +; SiO2 bodies 0; leaves spiral; flowers moderate in size; P = K + C, A basifixed, pollen more or less spiny, micropyle bistomal, style +; K persistent in fruit; seed operculum ["embryostega"] +, endostomal; embryo undifferentiated.
MAYACACEAE> Kunth - Stems ± prostrate; leaves apically bidentate, univeined, without a distinct sheath, uniseriate colleters +; flower single, apparently axillary; A 3, opposite K, anther dehiscence apical, ± porose, exothecium +, endothecium unthickened, ovules atropous. - 1/4-10. Mostly tropical and American (inc. S.E. U.S.A.), 1 sp. from W. Africa.
Eriocaulaceae + Xyridaceae: rosette plants; vessel elements with simple perforations; inflorescence usu. axillary, scapose, capitate, with involucral bracts; C clawed, A adnate and opposite C, exothecium +.
ERIOCAULACEAE Martynov - Flowers imperfect; inflorescence receptacle ± flat; pollen spiraperturate, 1 pendulous atropous ovule/carpel, micropyle endostomal. - 10/1160. Pantropical (to temperate).
1. Eriocauloideae - Plants usu. of aquatic habitats; roots and leaves with aerenchyma; C free, with black tips, staminate flowers: A adnate to C. - 1(-2?)/420. Pantropical (to Temperate).
2. Paepalanthoideae - C often connate, carpellate flowers: stylar appendages +, stigmas commissural. - 9/760. New World, but esp. tropical South America.
XYRIDACEAE C. Agardh - Vascular bundles amphivasal; median K membranous, deciduous. - 5/260. N. South America, Guyana Highlands in particular; pantropical to warm temperate.
1. Xyridoideae - Leaves distichous, equitant, isobifacial, ligulate; exothecium +, endothecium unthickened, staminodia 3, branched and with moniliform hairs on branch ends, placentation (intrusive) parietal. - 1/225-300. Pantropical to warm temperate, esp. Brasil.
2. Abolboidoideae - Tapetum amoeboid, pollen spherical, inaperturate, style solid; endosperm helobial. - 4/26. W. South America, Guyana Highlands in particular.
[[Anarthriaceae [Restionaceae + Centrolepidaceae]] [Flagellariaceae [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]: plant rhizomatous; flavones +; primary cell wall also with(1-3,1-4)-ß-D-glucans; sieve tube plastids with cuneate and other less densely packed crystals; leaves 2-ranked, with sheath; flowers small, imperfect [position unclear], T undifferentiated, membranous, endothecial wall thickenings girdle-like, pollen scrobiculate [minute pores penetrating tectum and foot layer], monoporate, annulate ["ulcerate"], 1 apical atropous ovule/carpel, micropyle bistomal, stigmas plumose, receptive cells on multicellular branches; embryo minute, undifferentiated, broad; deletions in ORF 2280 region.
Anarthriaceae [Centrolepidaceae + Restionaceae]: root hairs originating from any epidermal cells; chlorenchyma with peg cells; plant dioecious; A 3, opposite inner P, dorsifixed.
ANARTHRIACEAE D. F. Cutler & Airy Shaw - ORF 2280 +. - 3/11. West Australia.
Centrolepidaceae + Restionaceae: anthers bisporangiate/unithecal, embryo sac with compound starch grains, cells of nucellar epidermis anticlinally elongated.
CENTROLEPIDACEAE Endlicher - Inflorescence scapose; P 0; staminate flowers: A 1; carpellate flowers: G [5]. - 3/35. Hainan, IndoChina and Malesia to New Zealand, S. South America.
RESTIONACEAE R. Brown - Leaves lacking blades. - 58/520. Africa (inc. Madagascar), esp. the S.W, Hainan, IndoChina, Malay Peninsula, S. New Guinea to to New Zealand, esp. S.W. Australia, Chile.
Flagellariaceae [[Joinvilleaceae + Ecdeiocoleaceae] Poaceae]: trichoblasts from distal cell of pair; leaf blade with cross veins, ligule +; inflorescence paniculate, branches with adaxial swellings; fruit indehiscent; cotyledon not photosynthetic.
FLAGELLARIACEAE Dumortier - Climbers; stem dichotomising; leaf blade terminating in a tendril; ovule crassinucellate, micropyle endostomal, embryo sac bisporic [Allium-type]; ORF 2280 +. - 1/4. Paleotropics, to the Pacific Islands.
[Joinvilleaceae + Ecdeiocoleaceae] Poaceae: silica bodies cubic; epidermis with microhairs; foliar epidermis with long and short cells [latter silica-containing]; guard cells dumbbell-shaped; fusoid cells [large colorless cells in central mesophyll] +; stem hollow [level?]; nucellar cap +; first seedling leaf lacking lamina [possible]; 28 and 6.4 kb chloroplast genome inversion +.
Flagellariaceae + Ecdeicoleaceae: ?
JOINVILLEACEAE Tomlinson & A. C. Smith - Flowers perfect; embryo sac bisporic; rps14 gene to nucleus, pseudogene remaining in mitochondrion. - 1/2. Malay Peninsula to the Pacific.
ECDEIOCOLEACEAE D. F. Cutler & Airy Shaw - Rhizome with hairs; leaves lacking a blade; two adaxial outer T distinct, abaxial smaller; staminate flowers: pollen with operculum, wall without scrobiculi, with intraexinous channels. - 2/3. W. Australia.
POACEAE Barnhart - Sieve tube plastids also with rod-shaped protein bodies; leaves pseudopetiolate, midrib +, ligulate; two adaxial outer T distinct, abaxial smaller; staminate flowers: pollen with operculum, wall without scrobiculi, with intraexinous channels; carpellate flowers: G with 1 central crassinucellate ovule; fruit an achene, the testa closely adherent to pericarp, hilum long; peripheral layer of endosperm meristematic, embryo lateral, long, well differentiated, plumule lateral, cotyledon = absorbtive scutellum; collar [epiblast, the ligule of the cotyledon] conspicuous; trnT inversion in chloroplast genome, rps14 gene to nucleus, pseudogene remaining in mitochondrion. - 668/10025. Worldwide.
1. Anomochlooideae Potzdal - 2/4. Brasil, forests.
Pharoideae + Puelioideae + PACCMAD + BEP clades: inflorescence of laterally compressed, racemose, pedunculate spikelets, units with two sterile basal bracts [= glumes, spikelet bract + prophyll], flowers 2-ranked, each with lemma [?bract] and palea [?= connate adaxial tepals of the outer whorl]; lodicules [inner whorl tepals] 3; n = 12; 1 bp deletion in the 3' end of the mat K gene.
2. Pharoideae (Stapf) L. G. Clark & Judziewicz - Microhairs 0; leaves resupinate, lateral veins oblique; spikelet 1-flowered. - 3/14. Pantropical, in forests.
Puelioideae + PACCMAD + BEP clades [the bistigmatic clade]: spikelets disarticulate above the glumes; stigmas 2, two orders of stigmatic branching; 15bp ndhF insertion.
3. Puelioideae L. G. Clark, M. Kobay., S. Mathews, Spangler & E. A. Kellogg - 2/14. Tropical Africa.
PACCMAD + BEP clades: arm and fusoid cells 0; foliar cross veins 0; pseudopetiole 0; A 3, lodicules 2, stylodia free; 15 bp insertion in ndhF gene, disease resistance by the Hm 1 gene. Mostly non-forest.
Panicoideae + Arundinoideae + Centothecoideae + Chloridoideae + Micrairoideae Aristidoideae + Danthonioideae [PACCMAD clade]: phytoliths dumbbell-shaped; mesocotyl internode elongated, epiblast 0.
Panicoideae + Arundinoideae + Centothecoideae + Chloridoideae: 6 bp insertion in the 3' end of the mat K gene.
Panicoideae + Centothecoideae: hilum non-linear [or for a PACC clade].
4. Panicoideae Link - Culms usually solid; spikelets dorsally compressed, rachilla 0, 2-flowered, lower flower staminate or sterile [gynoecial cell death caused by Tasselseed2], one carpellate floret, spikelet dispersed as a 1-seeded unit by disarticulation below the glumes; starch grains simple; 5 bp insertion in the rpl16 intron, rps14 gene pseudogene lost; n = 5 (7) 9, 10 (12, 14); germination flap +. - 206/3270. Tropics to temperate.
5. Centothecoideae Soderstrom - Mesophyll differentiated into palisade and spongy mesophyll; chlorenchyma cells lobed [cf. arm cells]. - 12/32. Warm temperate to tropical forests. Poorly understood.
Micrairoideae + Arundinoideae + Chloridoideae + Aristidoideae + Danthonioideae: ligule hairy; lemma awned; starch grains compound.
Arundinoideae + Chloridoideae: hilum short.
6. Arundinoideae Burmeister - 14/33-38. Temperate to tropical, hydrophytic to xerophytic.
7. Chloridoideae Beilschmied - Microhairs with inflated distal cells; spikelets disarticulating above the glumes; 4 bp insertion in the rpl16 intron. - /1400. Tropical to warm temperate, more or less dry environments especially in Africa and Australia.
8. Micrairoideae Pilger - 8/170. Tropics.
Aristidoideae + Danthonioideae: awn trifid, or 3 awns.
9. Aristidoideae Caro - 3/385. Warm temperate, few in Europe.
10. Danthonioideae Barker & Linder - Haustorial synergid cells. - 19/250. Mesic to xeric.
Bambusoideae + Ehrhartoideae + Pooideae [BEP clade]: x = 12.
11. Bambusoideae Luersson - Woody; (mesophyll differentiated into palisade and spongy tissues); fusoid cells and strongly asymmetrically invaginated arm cells +; leaves pseudopetiolate, culm leaves often very different from the others. 84-101/940-1320. Tropical, often in forests.
12. Ehrhartoideae Link - 17/120. Widespread, esp. S. hemisphere.
13. Pooideae Bentham - Fructose oligosaccharides in stem; microhairs 0; stomata subsidiary cells with parallel sides; primary inflorescence branches 2-ranked; lemma usually with 5 nerves; styles joined only at the very base. - /3300. Largely N. temperate.
COMMELINALES + ZINGIBERALES: inflorescences with many-flowered cincinnal [helicoid cyme] branches; tapetum invasive or plasmodial.
COMMELINALES Dumortier
Vessel elements scalariform; seed coat testal and tegmic; endosperm abundant, helobial, cell wall formation in small chalazal chamber precedes that in large micropylar chamber. - 5 families, 68 genera, 812 species.
Commelinaceae + Hanguanaceae: cotyledon not photosynthetic.
COMMELINACEAE Mirbel - T = K + C, septal nectaries 0. - 40/652. Tropical and temperate.
1. Cartonematoideae (Pichon) G. Tucker - SiO2 bodies 0; flowers sessile, yellow; most of testa sloughed off. - 2/12. Mostly Australian, Triceratella Zimbabwe.
2. Commelinoideae - Cyanidin 3,7,3'-triglucoside +; stem collenchyma +; stem with narrow cortex and endodermis-like sheath enclosing vascular bundles that connect only at the nodes; raphide canals between veins; 3-celled glandular microhairs +; seed operculate. - 38/640. Tropical, also temperate, not Europe.
HANGUANACEAE Airy Shaw - Leaves with petiole, midrib and cross veins; plant dioecious; inflorescence branched-spicate, flowers sessile; staminate flowers: pollen inaperturate, exine spinulose; carpellate flowers; staminodes nectar-secreting, intra-ovarian trichomes +, 1 basal atropous tenuinucellate ovule/carpel; fruit a 1-seeded berry; seed bowl-shaped [placenta inside]; mesotesta and endotesta sclerified. - 1/6. Sri Lanka, South East Asia to Palau and N. Australia.
Philydraceae [Haemodoraceae + Pontederiaceae]: SiO2 bodies 0; styloids +; T with tannin cells, sclereids in placentae; T persistent in fruit.
PHILYDRACEAE Link - Inflorescence racemose; perianth petaloid, two-merous, outer whorl usually much larger than the inner, A 1, septal nectaries 0; exotesta with thick cellulose walls, endotegmen tanniniferous, seed operculum +. - 4/5. Australia (all genera) to Southeast Asia.
Haemodoraceae + Pontederiaceae: phenylphenalenones +; ektexine not tectate or columellate, endothecial cells with base-plates.
HAEMODORACEAE R. Brown - Arylphenalenones +; leaves equitant, isobifacial; cyme [usu.] bifurcated; exine (1-)2(-3)-layered [no foot layer], G inferior, ovules atropous, placentae swollen. - 14/116. Tropics and warm temperate regions.
2. Conostyloideae Lindley - Hairs branched; flowers enantiostylous, pollen porate, placentae with tanniniferous cells; cotyledon photosynthetic, hypocotyl +, primary root well developed. - 6/77. S.W. Australia.
PONTEDERIACEAE Kunth - Filaments hairy, pollen 2- or 3-sulcate. - 9/33. Tropics, also temperate, esp. New World.
ZINGIBERALES Grisebach
No aerial stem except when flowering; SiO2 in bundle sheath; sieve tube plastids also with starch grains; petiole bundles in arcs; cuticular waxes as aggregated rodlets; leaves with distinct petiole, midrib and fine cross venation; inflorescence bracts large, persistent; flowers large, monosymmetric; T = colored K surrounding C, A 5, adaxial member of inner whorl not developed, anthers long, pollen inaperturate, exine thin and spinulose to 0 [pollen not resistant to acetolysis], G inferior, outer integument 3< cells across, nucellar epidermal cells radially elongated, style long, stigma large, wet; fruit capsular; seeds arillate, operculate, operculum testal, micropylar collar [develops from outer integument and forms annular inpushing in perisperm surrounding operculum], endotesta sclerotised and silicified; endosperm nuclear, perisperm s.l. +, starchy, embryo plug-like; cotyledon not photosynthetic, ligulate, collar roots +. - 8 families, 92 genera, 2111 species.
MUSACEAE Jussieu - Plant monoecious; inflorescence bracts deciduous, floral bracts and bracteoles 0; 5 T similar, connate except adaxially, one [adaxial, inner whorl] free, short, concave. - 2/35. Africa, Himalayas to South East Asia, Philippines and N. Australia.
HELICONIACEAE Nakai - Inflorescence bracts coloured; T all similar, median outer T adaxial, ± free, other five connate, A 5, basally adnate to C, adaxial outer A staminodial, scale-like, 1 basal apotropous ovule/carpel; fruit a drupe; seed coat undifferentiated. - 1/100-200. Mostly tropical America, a few Celebes to the Pacific.
Strelitziaceae + Lowiaceae: petiole with adaxial and abaxial series of air canals; P whorls ± distinct, both coloured, adaxial A of inner whorl sterile, tapetum glandular [inc. Lowiaceae?], floral column [sterile apex of ovary] +, stigma 3-lobed; aril hairy.
STRELITZIACEAE Hutchinson - Inner lateral tepals postgenitally ± connate, forming labellum, inner adaxial tepal smaller, ± cucullate. - 3/7. Tropical South America, E. southern Africa, Madagascar.
LOWIACEAE Ridley - Cross veins in abaxial part of lamina; petiole long; inflorescence complex [repeating 1-flowered units, branching from bracts below the flower, the flower axillary]; flowers held upside down, K basally connate, functionally abaxial "petal" large, labellar, 2 adaxial "petals" small, enclosing A, nectary 0, outer integument 14-16 layers, stigma monosymmetric, dorsiventrally flattened, secretory tissue [viscidium] on adaxial side at base, lobes ± fimbriate; seed hairy, testa vascularized. - 1/15. S. China to Borneo.
Cannaceae + Marantaceae + Zingiberaceae + Costaceae: raphides 0; petiole with one series of air canals; guard cells with inner and outer ledges unequal; petiole short, poorly differentiated; C connate, both A whorls with two staminodes, adaxial A of inner whorl fertile; micropylar collar well developed, cells of exotesta longitudinally elongated; endosperm slight, chalazosperm +.
Cannaceae + Marantaceae: oblique cells in petiole; flowers asymmetrical; A ½, staminodes free, stigma not notably expanded.
CANNACEAE Jussieu - Mucilage canals in stem; style flattened; capsule muricate; operculum 0, mesotesta sclereidal, endotesta not sclereidal. - 1/19. New World (sub)tropics.
MARANTACEAE R. Brown - Lamina folding upwards at night, with S-shaped secondary veins, petiole well differentiated, with upper pulvinus; flowers in mirror-image pairs, pollination explosive, C A and style all basally fused, one inner staminode cucullate, another ± fleshy and with callosities, fertile half stamen often with a petaloid lateral appendage, 1 basal ovule/carpel, style under tension, becoming curved, pollen deposited on adaxial surface flat "stamp", secretory area adjacent. - 31/550. Tropics, esp. American, not in Australia.
Costaceae + Zingiberaceae: leaf ligulate; K connate, labellum from lateral staminodes of outer whorl and from the 2 staminodes of inner whorl, large, with narrow tube and distinct open limb, A 1, exine +, style slender, running between two half anthers, nectaries 2, on top of ovary, stigma cup- or funnel-shaped; chalazal mass in seed ± developed, hypocotyl well developed.
COSTACEAE Nakai - Aerial stem +; sheath with 1 series of adaxial air canals, no canals in petiole and lamina, vascular bundles adaxial; leaves spiromonostichous, sheath closed; inflorescence spicate-capitate; abaxial member of outer A whorl staminodial, all 5 staminodes connate, pollen aperturate, exine +; endosperm without starch; cotyledon blade-like, photosynthetic, with apical backwardly-directed process. - 4/110. Pantropical, esp. America and Papuasia-Australia.
ZINGIBERACEAE Martynov - Plant with ethereal oils; staminodes connate. - 46-52/1075-1300. (Sub)tropical, esp. South East Asia-Malesia.
1. Siphonochiloideae W. J. Kress - Rhizome fleshy, vertical. - 1/15. Africa and Madagascar.
2. Tamijioideae W. J. Kress - Rhizome fibrous; placentation parietal. - 1/1. Borneo.
3. Alpinioideae Link - Rhizome fleshy, plane of distichy parallel to the ground [position where?]; lateral staminodes of outer whorl