Summary of Apomorphies in /APweb/

Arrangement of Families and Orders, etc., in /APweb/

APG II System (see Angiosperm Phylogeny Group 2003)

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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a particular character, there may be disagreement over its interpretation (e.g. Austrobaileyales), or I have added unpublished information (e.g. Myrsinaceae and their relatives, nodal anatomy, etc.; Diapensiaceae, leaf ptyxis; Peridiscaceae, Centroplacaceae, etc.).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.


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.

GYMNOSPERMS  Back to Main Tree

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.


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 +.


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.


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; cyclopropyl amino acids +; 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.


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 four cellular [one-modular], stylulus short, hollow, stigma ± decurrent, dry; 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; pericycle with hippocrepiform sclereids; plant dioecious; hypanthium +, A sessile, vascular bundle branched towards apex, microsporogenesis successive, pollen ektexine granulate, 1 ovule/carpel, embryo sac monosporic, spore chalazal, embryo sac 9-nucleate, bipolar [two-modular]; 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.


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; tracheids with meshwork of coarse fibrils on their end walls; minute rhombic crystalls on stellate cells [astrosclereids, stellate parenchyma cells]; 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, stigma wet; exotesta palisade, anticlinal walls sinuous.

CABOMBACEAE A. Richard - Floating and rhizomatous; stem vascular tissue with two pairs of bundles; stomata anomocytic; flowers 3-merous, P members with single trace, A with slender filaments. - 2/6. World-wide.

NYMPHAEACEAE Salisbury - Vessel elements in roots, with extensive fibrillar network in the end plates; astrosclereids +; P members with sepaloid and petaloid areas. - 3/58. World-wide.

1. Nupharoideae Ito - Rhizomes stout, creeping; roots with well-developed pith; K 5-14, inner "tepals" small, nectary on abaxial surface, 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 [CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]: ethereal oils in spherical idioblasts [lamina and P ± pellucid-punctate]; tension wood 0; 12BP [4 amino acids] deletion in P1 gene.


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; testa vascularised, almost all walls thick, lignified; perisperm +. - 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, antipodal cells persisting; endosperm triploid.



?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, antipodal cells proliferating; 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; ovules lateral.


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 - Epidermal cells with single crystals; flowers with open development; P 3 [smaller] + 3 + 3, very thick, valvate, 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.


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, stylulus long, extragynoecial compitum +; 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 +, stigma sessile, stout, fleshy; 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 [Gomortegaceae + Atherospermataceae]] [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, ovule median, on early-initiated cross zone, vascular bundle not branching in chalaza.

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 1. - 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, archesporium single-celled, 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, style short, stigma stout. - 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; plants heterodichogamous; 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; exotesta palisade. - 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.


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, antipodal cells proliferating, 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.


Inflorescence a spadix, bract/bracteoles 0, with spathe; ovules staight [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, placentae apical, pendulous. - 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.


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 - Leaves two-ranked, margins minutely toothed; inflorescence complex, branched, spathe 0; 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: stem usu. aerial; styloids +; pseudopetiole apically geniculate; crystals often surrounding the embryo.

4. Pothoideae Engler - Secondary and tertiary veins reticulate; spathe not enclosing spadix. - 4/900. Tropical America, Madagascar to South and Southeast Asia, Malesia and N.E. Australia.

5. Monsteroideae - 12/360. Tropical South and Southeast Asia to the Pacific, South America (Africa).

6. 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).

Zamioculcadoideae + Aroideaee: plants monoecious (dioecious); spathe differentiated into tube plus blade, pollen extruded in strands. - 70/2670. Tropical and warm temperate.

7. Zamioculcadeae Bogner & Hesse - ? - 3: 21. Africa, Kenya to Natal.

8. Aroideae - 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 +.

Alismataceae [Butomaceae + Hydrocharitaceae]: 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.

ALISMATACEAE Ventenat - 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; fruit a follicle; embryo strongly curved, white; x = 7, 8. - 15/88. Pantropical, also temperate.

Hydrocharitaceae + Butomaceae: ?

BUTOMACEAE Mirbel - Leaves three-angled. - 1/1. Temperate Eurasia.

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.

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 + Juncaginaceae + Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: ?

APONOGETONACEAE J. Agardh - Inflorescence spicate, bracts 0. - 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.


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.




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 +; pollen orbicules with a circular perforation, ovary ± inferior in flower, ovules campylotropous; integuments lack cuticle; 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.


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 10-many, free, 1 (2) basal tenuinucellate ovule/carpel. - 8/48. Pantropical.

Stemonaceae + Pandanaceae + Cyclanthaceae: flowers other than 3-merous; placentation parietal, style 0.

STEMONACEAE Caruel - Distinctive alkaloids + [with a pyrrolo- or pyrido(1,2,-alpha) azepine nucleus]; 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 - Inflorescence with spathe, flowers in spirals; staminate flowers: T with abaxial gland, filaments swollen basally; carpellate flowers: 4(-8)-merous, staminodes opposite P, long-filiform; fruit baccate. - 11/225. Central and tropical South America.



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; T green, not spotted, ovules crassinucellate, styluli +; 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.



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).

Ixoliriaceae, etc. + Doryanthaceae + Iridaceae + Xeronemataceae + Hemerocallidaceae, etc., + Alliaceae, etc., + Asparagaceae, etc.: ?

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 [Iridaceae [Xeronemataceae [Hemerocallidaceae, etc. [Alliaceae, etc., + Asparagaceae, etc.]]]]: ?

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. - 66/2025. World-wide.

1. Isophysidoideae Takhtajan - 1/1. Tasmania.

Iridoideae [Patersonioideae [Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]]]: xanthone [mangiferin] +; inflorescence a rhipidium; (pollen operculate [often with two exine bands in a sulcus]), G inferior; endosperm 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.

Patersonioideae [Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]]: rhipidia 2, fused [binate], each unit with 2-many flowers; T connate, endothecium with base-plate or U-shaped thickenings; extra codon in rps4 gene.

3. Patersonioideae Goldblatt - Plant ± woody and rhizomatous; biflavonoids [amentoflavone] +; secondary growth +; flowers blue, last one day, filaments ± connate, pollen spherical, inaperturate, intectate; embryo small; two extra codons in rps4 gene. - 1/55. More or less open conditions, sub-Saharan Africa and Madagascar.

Geosiridoideae [Aristeoideae [Nivenioideae + Crocoideae]]: ?

4. Geosiridoideae Goldblatt & Manning - Echlorophyllous, saprophytic; leaves heterobifacial; flowers sessile; T connate basally only, microsporogenesis successive, nectary 0; seeds minute, dust-like; endosperm helobial, starchy. - 1/1: Geosiris. Madagascar, the Comores.

Aristeoideae [Nivenioideae + Crocoideae]: ?

5. Aristeoideae Vines - Plant rhizomatous; plumbagin +; flowers ± blue, usually last one day; T connate basally only; embryo small. - 1/55. More or less open conditions, sub-Saharan Africa and Madagascar.

Nivenioideae + Crocoideae: ?

6. Nivenioideae - Plant woody; secondary thickening +; unit of rhipidium with 1-2 flowers, flowers sessile; septal nectaries +; 1(-4) shield-shaped [tangentially flattened] seeds per loculus. - 3/14. Restricted to the S.W. Cape region, South Africa

6. Crocoideae G. T. Burnett - Leaf sheath closed; inflorescence spicate; binate rhipidium with a single flower, flowers sessile, obliquely monosymmetric (polysymmetric), endothecium with spiral thickenings, pollen aperture with one or a pair of longitudinal bands forming operculum, ovules campylotropous. - 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.).


Unlignified cell 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 Antilles

5. 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 +.

Joinvilleaceae + 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; chloroplast genome with trnT inversion, only 17 introns [those in clpP and rpoCI genes absent], 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: (benxoxazinoids, ergot alkaloids [latter synthesized by endophytes] +); 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.

Ehrhartoideae [Bambusoideae + Pooideae] [BEP clade]: x = 12.

11. Ehrhartoideae Link - 17/120. Widespread, esp. S. hemisphere.

Bambusoideae + Pooideae: characters?

12. 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.

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.


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.

Haemodoroideae Arnott - Roots red; A dimorphic, G secondarily superior, placental sclereids +. - 8/39. Tropics and warm temperate regions, not in southern S. America, only southern Africa, in S.E. Asia, not E. of Wallace's Line.

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.


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]: SiO2 bodies decorated [druse-shaped] and with troughs; raphides 0; petiole with one series of air canals; guard cells asymmetrical [with inner and outer ledges unequal]; petiole short, poorly differentiated; K = petalline K + C, C connate, both A whorls with two staminodes, adaxial A of inner whorl fertile, tapetum amoeboid, non-syncytial, micropyle endostomal, nucellar cells radially elongated {"nucellar pad"]; micropylar collar well developed, [annular wedge-shaped invagination into the nucellus formed by the inner integument]; cells of exotesta longitudinally elongated; chalazosperm [perisperm of some authors] +, 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; funicle hairy, imbibition lid on raphe, micropylar collar/operculum 0, exotesta and epidermis of chalaza a malpighian layer, mesotesta sclereidal, endotesta not sclereidal. - 1/10. 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 +, resistant to acetolysis, style slender, hollow, 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 very small or 0 [labellum formed by the two staminodes of the inner whorl alone]; endosperm without starch. - Indo-Malesia, tropical Australia; Renealmia, American and African tropics.

4. Zingiberoideae Hasskarl - Rhizomes fibrous; plane of distichy at right angles to the ground; lateral staminodes of outer whorl also free from labellum, labellum adnate to filament and forming tube [Globbeae]; endotesta parenchymatous; starch grains of perisperm compound. - Indo-Malesia, tropical Australia.



Aquatic herb; roots 0; vessels 0; nodes?; cuticle and stomata 0; leaves opposite, margins spiny-toothed; plant monoecious, P not vascularised, A extrorse, connective apically produced, tapetum amoeboid, pollen grains inaperturate, exine much reduced, tubes branched, G 1, postgenital fusion by secretion, with 1 atropous apical unitegmic ovule; seed coat ± obliterated; endosperm [with small initial micropylar cell], suspensor 0. - 1 family, 1 genus, 1-2+ species.

CERATOPHYLLACEAE Gray - One vegetative bud/node, flowers extraxillary, alternating with leaves; endosperm 0, embryo large, green. - 1/2 or more.  World-wide.

EUDICOTS: Root epidermis derived from root cap; stomata anomocytic; flowers (dimerous), cyclic, P members with a single trace, A (opposite P), few, filaments fairly slender, anthers basifixed, pollen tricolpate, G with complete postgenital fusion, style solid [?here]; seed coat?


Young stem with separate bundles; rays exclusively wide multiseriate [and in wood, where present]; nodes 3:3; sieve tube plastids large S-type; cuticle waxes as clustered tubules, nonacosan-10-ol the main wax; seed exotestal; endosperm development?, embryo size? - 7 families, 199 genera, 4445 species.

EUPTELEACEAE K. Wilhelm - P 0, anthers valvate, connective prolonged, carpels stipitate, ± ascidiate, stigma brush-like; fruit one-seeded, samaroid. - 1/2. Temperate South East Asia.

Papaveraceae [[Circeasteraceae + Lardizabalaceae] [Menispermaceae [Berberidaceae + Ranunculaceae]]]: vessel elements with simple perforation plates; leaves ternately compound or palmately lobed, articulated, and/or with palmate venation.

PAPAVERACEAE Jussieu - Herbaceous; roots diarch [lateral roots 4-ranked]; leaf base broad; P = K + C, K 2, median, distinct from C, small, C 4, G [2], placentation parietal, style +; capsule septicidal, (with persistent woody placenta); endotesta well developed. - 44/760. Largely N. Temperate, also S. Africa, scattered in South America, etc.

1. Pteridophylloideae - Venation pinnate; A 4, 1-2 ovules/carpel. - 1/1. Japan.

Papaveroideae + Fumarioideae: isoquinoline protopine +; laticifers, articulated or not, anastomosing or not; vessel elements with simple perforations; petiole bundles arcuate; leaves usu. soft, ± fleshy, micropyle bistomal; (seeds arillate); endotesta with calcium oxalate crystals; exotegmen fibrous or not, endotegmen walls thickened; endosperm nuclear.

2. Papaveroideae Eaton - Plant with latex; K enveloping flower in bud, fugaceous, C crumpled in bud, A many, style short/0. - 23/230. Largely N. temperate.

2A. Papavereae Dumortier - 8/95-125. N. (warm) temperate, also South America (Argemone) and S. Africa and Cape Verde Islands (1 sp. in each - Papaver).

2B. Chelidonieae Dumortier - 9/48. East Asia and E. North America (Europe, C. and S. America, West Indies).

2C. Eschscholtzieae Baillon - Pollen polycolpate; fruit dehiscing explosively from the base. - 3/16. W. North America.

2D. Platystemoneae Spach - Styles separate. - 3/5. W. North America.

3. Fumarioideae Eaton - Flowers strongly transversely disymmetrical, A 6, nectaries at base of stamens. - 20/530. Mostly N. temperate, also S. Africa.

3A. Hypecoeae Dumortier - Inner petals 3-lobed, A 4, two with two vascular strands [each 2 A connate], pollen dicolpate, deposited on inner petals; stylodia 2; seeds covered with rectangular crystals. - 1/18. Mediterranean to W. China.

3B. Fumarieae Dumortier - K minute, two outer C spurred, inner C apically coherent, with median joint, A in two groups of three, 2 anthers dithecal and 4 monothecal, pollen deposited on stigma; seed with chalazal aril; suspensor cells like a small bunch of grapes. - 19/505. Eurasia, North America, North and South Africa, mountains of E. Africa; ¾ species in Sino-Himalayan region.

[Circeasteraceae + Lardizabalaceae] [Menispermaceae [Berberidaceae + Ranunculaceae]]: rays broad; flowers 3-merous, K, C, and A, or P and A opposite each other, outer P members with three or more vascular traces.

Circeasteraceae + Lardizabalaceae: A extrorse.

CIRCAEASTERACEAE Hutchinson - Herbs; venation dichotomous; pollen tectum striate, carpels asxcidiate, ovules tenuinucellate. - 2/2. N. India to S.W. and W. China.

Circaeaster Maximowicz - Bracteoles 0; P 2-3, A latrorse?, anthers bisporangiate/monothecal, ovule straight [atropous]; n = 15. - 1/1.

Kingdonia Balfour f. & W. W. Smith - Annual; leaves two-ranked; P 5(-7), 8-13 clavate glands, G 3-9, ovules hemianatropous; n = 9. - 1/1.

LARDIZABALACEAE R. Brown - Vines/lianes; leaves compound; flowers imperfect; placenta fleshy in fruit. - 9/36. South East Asia and Chile.

1. Sargentodoxoideae Thorne & Reveal - Cork deep-seated; G ascidiate, many, 1 pendulous ovule/carpel. - 1/1. China.

2. Lardizabaloideae Kosteletzky - 8/35. South East Asia and Chile (Lardizabala, Boquila).

Menispermaceae [Berberidaceae + Ranunculaceae]: berberin + [a benzylisoquinoline - scraped stems yellow in color]; endosperm nuclear.

MENISPERMACEAE Jussieu - Lianes; serial cambia frequent; petiole pulvinate at apex and base, lamina entire; flowers imperfect, P with a single trace, anther thecae superposed, pollen with endoapertures [colporate], ovules 2/carpel, 1 aborting; fruit a drupe, with condyle [placentar intrusion]; embryo long, cotyledons ± foliaceous. - 70/420. Pantropical, usually lowland.

1. Tinosporoideae W. Wang & Z. D. Chen - Endosperm ruminate.

2. Menispermoideae W. Wang & Z. D. Chen - Style lateral to basal; endocarp often sculpted; (endosperm ruminate; 0), embryo curved, cotyledons narrow.

Berberidaceae + Ranunculaceae: rhizomatous herbs; roots diarch [lateral roots 4-ranked]; vascular bundles V-shaped, in herbaceous taxa often closed, scattered or in concentric rings; leaf base broad, (lateral petiolar stipules +); ± petaloid staminodial nectaries +, nucellar cap +, outer integument at least 4 cells thick; seed exotestal.

BERBERIDACEAE Jussieu - Flowers with cortical vascular system, 6 nectariferous "C", A 6, with flaps, G 1, ascidiate, postgenital occlusion by secretion, outer integument 5-11 cells thick, antipodals endopolypolypoid, stigma broad; exotesta lignified, with oblong-fibrous (cuboid) cells; endosperm with hemicellulose. - 14/701. Mostly East Asia and E. North America, also South America, N. Africa, general N. temperate.

1. Nandinoideae Heintze - Lowermost branch of inflorescence from axil of ± expanded leaf; G with 1-2 (4) ovules; fruit a berry or pericarp evanescent or bladder-like; funicle swollen; n = 8, 10. - 4/15. E. Europe to Japan.

2. Berberidoideae Kosteletzky - "C"/staminodia with paired basal nectaries, stamens sensitive, microsporogenesis successive [?all], pollen 6-12 colpate, or apertures irregular (spiraperturate), wall undifferentiated; fruit a berry; embryo long; n = 7. - 2/601. General N. temperate, also South America, N. Africa.

3. 3. Podophylloideae Eaton - Pollen striate; n = 6. - 8/75. Mostly (Europe to) East Asia (some desert xerophytes) and W. or E. North America.

RANUNCULACEAE Jussieu - P uniseriate, A many, spiral, G many, spiral. - 62/2525. ± World-wide, but esp. (N.) temperate, not lowland tropics.

Hydrastidoideae + Glaucidioideae: vessel elements with scalariform perforation plates; medullary bundles +; vascular bundles flat; petiole bundles annular and medullary; palisade mesophyll 0; leaves 2-ranked; flowers single, terminal; nectaries 0, outer integument 4-10 and inner integument 3-5 cells across, stigma bilobed; follicle dehiscing abaxially as well.

1. Hydrastidoideae Martynov - Petiole base on rhizome encircling stem; K 2-3, with a single trace, C 0, pollen tectum striate-reticulate, micropyle zig-zag; exotesta strongly palisade, exotegmen lignified, both multiplicative; x = 10. - 1/1. C. and E. North America.

2. Glaucidioideae Loconte - Berberin 0; flowers with cortical vascular system, A centrifugal, G 2, basally connate, many tenuinucellate ovules/carpel, archesporium multicellular, nucellar cap massive; seeds winged, outer integument vascularised; x = 13. - 1/1. Japan.

Coptoideae + Isopyroideae + Ranunculoideae: xylem surrounding phloem; petiole bundles with associated lignification; P 5-merous, "C" 0-13, usu. obviously nectariferous, very diverse in form.

3. Coptoideae Tamura - X = 8. - 3/17. East Asia, E. and W. North America.

Isopyroideae + Ranunculoideae: ovules apotropous [when single].

4. Isopyroideae Rafinesque - Trichomes clavate or saccate, lamina surface often papillate; x = 7. - 9/450. N. temperate, also South America, Africa and New Guinea.

5. Ranunculoideae Arnott - Berberin 0; stomata >35 µm long; trichomes pyriform; x = 8. - 46/2025. Worldwide, but not tropical.


SABIALES Takhtajan  

P = K + C, pollen colporate, nectary disc-like, carpels 2-3, connate, 1 or 2 pendulous campylotropous ovules/carpel, micropyle 0; seed with condyle [placental intrusion]; endosperm helobial, embryo long, curved. - 1 family, 3 genera, 100 species.

SABIACEAE Blume - 3/100. South East Asia to Malesia, tropical America.


PROTEALES Dumortier  

Epidermal waxes with tubules [2/3]; nodes?; leaf teeth?; stipules surrounding stem; 1-2 pendulous ovules/carpel, stigma dry; seed coat?; endosperm development?, slight or 0, embryo long. - 3 families, 82 genera, 1610 species.

NELUMBONACEAE Berchtold & J. Presl - Aquatic; articulated laticifers +; prophyll adaxial; leaves in groups of three along the stem, vertically 2-ranked, sheathing cataphyll on one side, then cataphyll and expanded leaf on the other side; leaf peltate, involute, main veins dichotomous, proceeding to margin; flowers axillary, large, with cortical vascular system; K 2, C 10-30, spiral, A many, spiral, connective with long appendage, receptacle massive, with emergent druses, carpels embedded in receptacle, outer integument ca 30 cells and inner integument 8-10 cells thick, nucellar cap and funicular obturator +, antipodals multiplying, multinucleate, stylodium 0; several leaf primordia, radicle aborting, roots adventitious. - 1/1-2. N. temperate.

Platanaceae + Proteaceae: woody; myricetin +, benzylisoquinoline alkaloids 0; wood with broad rays [8+-seriate]; flowers 4-merous ; A = and opposite P, carpels with 5 vascular bundles, hairy, postgenital fusion complete, ovules atropous, stylodium long; endosperm nuclear.

PLATANACEAE T. Lestibudois - Hairs branched; plant monoecious; inflorescence capitate; 2 ovules/carpel; fruit an achene with tuft of hair at the base. - 1/10. North Temperate.

PROTEACEAE Jussieu - Lateral roots of limited growth, forming clusters [proteoid roots]; hairs with 2 short cells, one in epidermis, apical cell elongated; stipules 0; bracteoles 0; P decussate, valvate, pollen triangular in polar view, oblate, porate, pores broadly operculate, arranged according to Garside's Rule, G 1, micropyle bistomal. - 80/1600. Largely southern hemisphere, esp. Australia and S. Africa.

1. Bellendenoideae P. H. Weston - Al accumulation +; bracts 0; 2 ovules/carpel; fruit dry, indehiscent; endosperm slight; n = 5. - 1/1. N.E. Australia.

Persoonioideae + Grevilleoideae + Symphionematoideae + Proteoideae: stomata brachyparacytic; P connate, A adnate to P, more or less sessile, 4 nectary lobes +; endosperm +.

2. Persoonioideae - Proteoid roots 0; cotyledons obreniform; chromosomes 13-26 µm long. - 5/110. Mostly Australia, also New Caledonia and New Zealand.

Grevilleoideae + Symphionematoideae + Proteoideae: flowers vertically or obliquely monosymmetric [P split to base on one side, or 3 P connate, 1 free]; x = 14, chromosomes 0.5-5 µm long (mean).

3. Grevilleoideae Engler - Al accumulation +; sieve elements with rosette-like non-dispersive protein bodies; paired flowers +. - 45/855. Australia, to Southeast Asia, South America, Africa and Madagascar.

Symphionematoideae + Proteoideae: fruit indehiscent.

4. Symphionematoideae P. H. Weston & N. P. Barker - Proteoid roots 0; nectaries 0. - 2/3. S.E. Australia, inc. Tasmania.

5. Proteoideae - Al accumulation infrequent; sieve elements with non-dispersive protein bodies; flowers sessile; fruit single-seeded. - 25/640. S. Africa, Australia.

TROCHODENDRALES + BUXALES + CORE EUDICOTS: rps 11 mitochondrial gene lost.


Nodes 3:3; idioblasts in cortex; stomata laterocytic; leaves toothed, secondary veins proceed to a clear, persistent cap, lateral veins also enter; bracteoles 0; filaments thin, anthers valvate, latrorse, G laterally connate, nectariferous abaxially, with 5 vascular bundles, ovules apotropous, integuments long, micropyle bistomal; fruit an aggregate of follicles, stylodia becoming ± basal on the outer surface; seeds flattened, with marginal flange, chalazal protrusion and hair-pin bundle, exotegmic cells thick-walled, tracheidal; endosperm development? - 1 family, 2 genera, 2 species.

TROCHODENDRACEAE Eichler - 2/2. East Asia.

BUXALES + CORE EUDICOTS: rps2 mitochondrial gene lost.

BUXALES Reveal  

Steroidal pregnan pseudoalkaloids +; true tracheids + ; stomata cyclocytic; lamina margin entire; inflorescence racemose, flowers small, imperfect; anthers ± dorsifixed, micropyle bistomal, stylodia stigmatic their entire length, with two crests; seed testal, with several cell layers; endosperm development?, embryo size? - 2 families, 5 genera, 72 species.

DIDYMELACEAE Leandri - P a scale [carpellate flower] or 0, A extrorse, G 1, ovule single; fruit a drupe. - 1/2. E. Madagascar.

BUXACEAE Dumortier - Stomata cyclocytic. - 4/70. ± World-wide, not Arctic.

1. Sarcococceae - Pollen pantoporate, with crotonoid exine processes; fruit a drupe. - 3/22. E. Asia to W. Malesia, E. North America, N. South America.

2. Buxeae - Fruit an explosively-dehiscent capsule. - 1/30. Scattered, inc. Madagasacar, only Central America and West Indies in New World, not Australia nor much of Malesia.

CORE EUDICOTS: ellagic and gallic acids common, cyanogenesis via phenylalanine, isoleucine or valine pathways; micropyle?; AP3 duplication [euAP3 + TM6 paralogs], PI-dB motif +, small deletion in the 18S ribosomal DNA common.


Ellagic acid +; vessel elements?; sieve tube plastids with protein crystalloids and starch; pith with sclerenchymatous diagrams; leaves with toothed margins, 2ndary veins palmate; plants dioecious, flowers small; P 0; seed coat?; endosperm cellular. - 2 families, 2 genera, ca 45 species.

GUNNERACEAE Meisner - Herbs; Nostoc colonies in stem and root; nodes multilacunar; glands on stem, colleters +; flowers ebracteolate, G [2], inferior, uni(bi)locular, 1 (2) apical ovules, embryo sac tetrasporic 16-celled [Peperomia-type]. - 1/40-50. Circum S. Pacific, Africa, Madagascar.

MYROTHAMNACEAE Niedenzu - Aromatic-resinous shrubs; plant glabrous; nodes split-lateral; leaves opposite, basally forming a sheath, plicate, stipules 2, small, persisting on the petiolar sheath; G with surface oil cells and 5 vascular bundles, embryo sac bisporic 8-celled [Allium-type]. - 1/2. Africa and Madagascar.

ROSIDS + ASTERIDS, ETC.: root apical meristem closed; flowers rather stereotyped: 5-merous, K and C distinct, A = 2x K, (often numerous, but then usually fasciculate and/or centrifugal), pollen tricolporate, (nectary disc +), [G 5], [3] also common, compitum +, placentation axile, stigma not decurrent; endosperm nuclear; fruit dry, dehiscent, loculicidal [when a capsule].

The sepals have three traces, the petals have one; taxa with numerous stamens, often developing from 5(10) primordia and/or centrifugal, are found in most orders.

DILLENIALES + CARYOPHYLLALES: tension wood 0; successive cambia +; vessel elements with simple perforations; wood with SiO2 bodies; nodes 3 or more:3 or more; leaves spiral; K persistent in fruit.

Note: for Caryophyllales, see just before asterids. The above might be synapomorphies if Dilleniales are indeed sister to Caryophyllales.

DILLENIALES Hutchinson  

Secondary veins proceeding straight to the teeth; A many, G separate, ?micropyle; fruit a follicle, seed arillate; endotesta ± palisade, lignified, exotegmen usu. tracheidal. - 1 family, 10 genera, 300 species.

DILLENIACEAE Salisbury - C crumpled in bud; K and filaments persistent in fruit. - 10/300. Tropical and warm temperate.

Saxifragales [Vitales + Rosids]: Stipules +; nodes 3:3.


Ellagic acid +; anthers basifixed, with basal pit, anthers transversely sagittate, ?pollen, floral apex flat-concave early in development [G often (semi-)inferior], carpels free, at least apically, micropyle bistomal, stigmas decurrent; fruit dry; seeds ± exotestal; endosperm type?, embryo size? - 16 families, 112 genera, 2470 species.

PERIDISCACEAE Kuhlmann - Leaves entire; G 1-locular, with 6-8 pendulous apical ovules; seed 1, coat "tanniniferous", walls thin, ± collapsed; endosperm copious, cell walls thick, pitted. - 3/9. South America, tropical W. Africa.

[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae + Daphniphyllaceae]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae [Haloragaceae + Penthoraceae]]]] [[Iteaceae + Pterostemonaceae] [Grossulariaceae + Saxifragaceae]]]]: ?

[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae + Daphniphyllaceae]]]: mitochondrial coxII.i3 intron 0.

PAEONIACEAE Rafinesque - Leaves ternately compound; flowers large, P spiral, stamens many; testa fleshy, vascularised, exotestal cells palisade, variously thickened, the hypodermis palisade, ± lignified; early embryo coenocytic. - 1/33. N. Temperate, especially East Asia.

[Altingiaceae [Hamamelidaceae [Cercidiphyllaceae + Daphniphyllaceae]]]: cuticle waxes as tubules, nonacosan-10-ol the main wax; inflorescence racemose, flowers sessile; anthers valvate, with protruding connectives; embryo long.

ALTINGIACEAE Horaninow - Iridoids +; secretory canals with aromatic contents; stipules petiolar; inflorescence ± capitate; P 0; staminate flowers: pollen polyporate; carpellate flowers: intercarpellary protrusions, etc., present [representing sterile flowers]; seed ?exotegmic. - 1/13. E. Mediterranean, East Asia to Malesia, Central America.

[Hamamelidaceae [Cercidiphyllaceae + Daphniphyllaceae]]]: pollen colpate.

HAMAMELIDACEAE R. Brown - Hairs stellate; C ribbon-like, adaxially circinate. - 27/82. Tropical to temperate, esp. East Asia to Australia, not South America.

1. Exbucklandoideae Harms - Inflorescence capitate. - 3/4-14. East Asia, to Assam, East Malesia, to Sumatra.

2. Disanthoideae Harms - Anthers with longitudinal slits. - 1/1. E. China, Japan.

3. Hamamelidoideae Harms - Leaves with craspedodromous venation; anthers with flaps, 1 fertile ovule/carpel; fruit with ballistic dispersal of seeds. - 23/78. Tropical to temperate, esp. East Asia to Australia, not South America.

Cercidiphyllaceae + Daphniphyllaceae: plant dioecious, inflorescence racemose; flowers small, C 0; pistillate flowers: G superior.

CERCIDIPHYLLACEAE Engler - Prophyll single, adaxial; leaves [of long shoot] opposite, stipule adaxial-petiolar; plant dioecious; inflorescence capitate, flower a pseudanthium, bracteoles 0; P 0, carpellate flowers: G 1, with abaxial suture, stigma long-decurrent; seeds winged. - 1/2. China and Japan.

DAPHNIPHYLLACEAE Müller Argoviensis - Iridoids +; leaves entire, venation pinnate, stipules 0; flowers pedicellate; staminate flower: filament with three traces; embryo short. - 1/10. East Asia to Malesia.

[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae [Haloragaceae + Penthoraceae]]]] [[Iteaceae + Pterostemonaceae] [Grossulariaceae + Saxifragaceae]]: petiole bundle(s) arcuate; cuticle waxes not tubular; ovules apotropous [all?]; K persistent, withered; endosperm cellular.

Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae [Haloragaceae + Penthoraceae]]]: stem with endodermis; nodes 1:1; stipules 0.

CRASSULACEAE Jaume Saint-Hilaire - ± herbaceous leaf succulents; CAM metabolism; sedoheptulose [sugar reserve] +; sieve tube plastids lacking starch grains; rays 0; lamina with hydathodes; anthers median sagittate, G = C, free, with nectariferous scales or flaps at the base of each carpel; endosperm chalazal haustorium +, embryo long, basal cell of suspensor with mycelium-like haustorial branches; x = 8. - 34/1370. ± Cosmopolitan, esp. the Cape region and Mexico, but few in S. South America and Australia, not in Polynesia, frequently in drier regions.

Crassuloideae Burnett - Leaves opposite, with several marginal hydathodes; A = K, slightly introrse, ovules tenuinucellate; follicles releasing seeds through apical pore; exotesta with sinuous anticlinal walls, unipapillate; first division of micropylar endosperm cell in horizontal plane. - 2/196. Southern Africa, to S.W. Arabia, "Tillaea" more or less world-wide, the only Crassulaceae in Australia.

Kalanchoideae + Sempervivoideae: lamina with (sub)apical hydathode; seeds costate; first division of micropylar endosperm cell in vertical plane.

Kalanchoideae A. Berger - Plant ± woody; crystal sand +; C connate, A with spherical connective prolongation; seeds 4-6-costate, with a micropylar corona; x = 9. - 4/200. Old World, Especially the Karoo in southern Africa, but extends to South East Asia and Malesia,

Sempervivoideae Arnott - Seeds >6-costate; suspensor with vertical division in micropylar cell. - 28/975. Largely N. hemisphere.

Aphanopetalaceae [Tetracarpaeaceae [Haloragaceae + Penthoraceae]]: ?

APHANOPETALACEAE Doweld - C 0, pollen with rugulate-stellate surface, one apical ovule/carpel; fruit a one-seeded nut, calyx enlarged, spreading. - 1/2. W. and E. Australia.

Tetracarpaeaceae [Haloragaceae + Penthoraceae] [= Haloragaceae s.l.]: ?

TETRACARPAEACEAE Nakai - Fibrous endothecium 0, nectary 0, G superior, free, stigma sessile. - 1/1. Tasmania.

Penthoraceae + Haloragaceae: Herbs.

PENTHORACEAE Britton - Fruits circumscissile at the base of the free portion of each carpel. - 1/1-3. East and South East Asia, E. North America.

HALORAGACEAE R. Brown - Plant monoecious; stylodia with swollen bases; fruit nut-like; haustorial embryo suspensor +. - 8/145. World-wide, especially Australia.

[Iteaceae + Pterostemonaceae] [Grossulariaceae + Saxifragaceae]: leaves spiral; hypanthium +, A = and opposite K; fruit septicidal.

Iteaceae + Pterostemonaceae: [= Iteaceae s.l.] C-glycosylflavones +; stipules +; placentation axile, micropyle?, styluli long; endosperm sparse.

ITEACEAE J. Agardh - Flavonols 0; pith chambered; pollen bilateral, 2-porate, ektexine homogeneous. - 2/18. South East Asia to W. Malesia, E. North America, E. and S. Africa.

PTEROSTEMONACEAE Small - A 5, opposite K, filaments flattened, toothed, 5 staminodes opposite C. - 1/3. Mexico.

Grossulariaceae + Saxifragaceae: G [2-3].

GROSSULARIACEAE de Candolle - C small, pollen 5-15-porate, with distinctively rugose ectoapertures, G inferior, placentation parietal, style single, long; fruit baccate; seeds hard, arillate, exotestal cells palisade, mucilaginous. - 1/150. Temperate N. hemisphere, also along the Andes.

SAXIFRAGACEAE Jussieu - Herbs. - Ca 33/540. Mostly N. temperate and Arctic (S. temperate, tropical mountains).

CYNOMORIACEAE Lindley - Echlorophyllous root parasite; plant monoecious; staminate flowers: A 1, adnate to P, nectary-stylodium +; carpellate flowers: G 1, inferior, 1 pendulous atropous tenuinucellate unitegmic ovule, integument ca 7 cells thick, style long; fruit a nut; testa ca 7 cells across, persistent, cells little thickened; endosperm copious, thick-walled, embryo undifferentiated. - 1/2. Mediterranean to C. Asia.

VITALES + ROSIDS: Anthers ± dorsifixed, transition to filament narrow, connective thin.

VITALES Reveal  

Nodes 3-7:3-7; sieve tube plastids with protein crystalloids and starch; raphide bundles +; pearl glands + [spherical, multicellular glands with an apical stoma]; leaves with glandular teeth; common stamen-petal primordia, A = and opposite C, 2 apotropous ovules/carpel, micropyle?, style +, short; fruit a berry, K deciduous; seeds perichalazal, ± ruminate, testa multiplicative, exotesta fleshy, endotesta several-layered, lignified, crystalliferous, exotegmen (crossed) tracheidal. - 1 family, 14 genera, 850 species

VITACEAE A.-L. de Jussieu - 14/850. Pantropical and (warm) temperate.

1. Leeoideae Burmeister

- Raphides barbed; stipules borne along petiole margin, sheathing; C basally connate, A adnate to C, connate, with a lobed tube, nectary disc 0, G [3 (4)], loculi divided. - 1/24. Most Indo-Malesian, few Africa and Madagascar.

2. Viticoideae Eaton

- Climbers; raphides smooth; tendrils opposite the leaves; G 2. - 13/825. Pantropical and (warm) temperate.

ROSIDS: embryo long; chloroplast infA gene defunct, mitochondrial coxII.i3 intron 0.

EUROSID I: endosperm scanty.


Harman alkaloids +; style +; seeds ± exotestal; endosperm 0; chloroplast infA gene +. 2 families, 27 genera, 305 species.

KRAMERIACEAE Dumortier - Hemiparasitic; flowers monosymmetric; stipules 0; K petaloid internally, abaxial larger than the others, (2) 3 adaxial C clawed, ± connate, 2 abaxial smaller, not clawed, glandular [often secreting lipid], A (3) 4, anthers porose, endothecial cells with thickening parallel to long axis of cells; G [2], adaxial member much reduced, 2 collateral pendulous ovules/carpel, stigma small, recessed; fruit nut-like, with retrorsely barbed spines; seed 1, exotestal cells enlarged, tanniniferous; cotyledons cordate/auriculate at the base. - 1/18. S.W. U.S.A. to Chile, the West Indies.

ZYGOPHYLLACEAE R. Brown - Leaves opposite, compound, no terminal leaflet. - 22/285. Dry and warm temperate, also tropical.



Stomata ?; leaf teeth?, stipules +; inflorescence cymose; flowers small, micropyle bistomal; chloroplast infA gene +. - 3 families, 93 genera, 1355 species.

LEPIDOBOTRYACEAE J. Léonard - Cristarque cells in bundle sheath; stamens paracytic; leaves 2-ranked, articulated with petiole, entire, stipel single, long; plant dioecious, inflorescences terminal, congested; K and C similar in size, A 10, of two lengths, ± connate basally, anthers basifixed, nectary on inside of staminal tube, G [2-3], 2 collateral apical epitropous, pachychalazal ovules/carpel; fruit a septicidal capsule, endocarp distinct, columella persisting. - 2/2-3. West Africa, Central and South America.

Celastraceae + Parnassiaceae: hexitol dulcitol +, ellagic acid 0; nodes 1:1; leaves with veins running to congested deciduous tooth; A = C, G opposite C, ovules ± tenuinucellate, stigmas commissural.

PARNASSIACEAE Martynov - Herbs; young stem with separate bundles; staminodes opposite C, nectar at base, placentation parietal, stigmas commissural. - 2/51. N. temperate to Arctic, Mexico, Chile.

CELASTRACEAE R. Brown - 89/1300. Largely tropical, but also temperate.



HUACEAE A. Chevalier - Hairs stellate or peltate; cristarque cells +; stomata paracytic; leaves with basal glands on margin or abaxial surface; inflorescence fasciculate; G [5], unilocular, with basal ovules; seed 1, testa with vascular bundles, exotegmen of lignified palisade cells. - 2/3. Tropical Africa.

Vessel element type?; mucilage cells +; stomata ?; leaves compound, with terminal leaflet; micropyle bistomal, stylodia +; integument multiplicative, endotesta crystalliferous and palisade, exotegmen fibrous/tracheidal. - 6 families, 58 genera, 1810 species.

Connaraceae + Oxalidaceae: plant construction sympodial; benzoquinone rapanone +, ellagic acid 0; roots diarch [lateral roots 4-ranked]; wood rays uniseriate; sieve tube plastids with protein crystalloids; druses 0; cuticle wax platelets as rosettes; leaflets articulated, margins entire, stipules 0; C postgenitally basally united, with uniseriate glandular hairs, nectary extrastaminal, A of two whorls of different lengths, with uniseriate glandular hairs; exotesta ± fleshy.

CONNARACEAE R. Brown - Lianes; hairs uniseriate; wood commonly siliceous or with SiO2 grains; K connate; G free, 2 collateral near-basal ovules/carpel; fruit a follicle; seed 1, testa black, vascularised, sarcotesta +. - 12/180. Pantropical, especially Old World.

OXALIDACEAE R. Brown - Juice acrid, with oxalates; C contorted, often clawed, A extrorse, ovules tenuinucellate, both integuments at least 3 cells thick, stigmas spathulate/capitate; fruit a ± ribbed/angled capsule or berry; seed often with mucilaginous testa, explosive; endosperm frequent, starchy.

6/770. Usu. tropical (esp. at high elevations) or subtropical.

Cunoniaceae [[Brunelliaceae + Cephalotaceae] Elaeocarpaceae]: K valvate, postgenitally coherent by hairs.

CUNONIACEAE R. Brown - Leaves opposite, stipules interpetiolar; filaments incurved in bud, longer than petals, pollen dicolpate; fruit a septicidal capsule; endosperm starchy. - 26/280. Largely temperate and tropical S. hemisphere, few African.

[Brunelliaceae + Cephalotaceae] Elaeocarpaceae: inner integument 3-5 cells across.

Brunelliaceae + Cephalotaceae: P uniseriate, G free, 2 basal ovules/carpel, stylodia recurved, stigma decurrent; fruit a follicle.

BRUNELLIACEAE Engler - Leaves opposite, leaflets stipellate, 2ndaries prominent, proceeding to the margin, stipules cauline; inflorescence cymose; flowers small; endocarp separating from the rest, K persistent; seeds shiny, raphe ± aril-like, coat with subepidermal sclerenchymatous layer and palisade innermost layer; endosperm mealy. - 1/55. Central and South America and the Antilles.

CEPHALOTACEAE Dumortier - Insectivorous herbs; leaves simple, some ascidiate and with a lid, margins entire, stipules 0; inflorescence scapose, racemose, branches dichasial; flowers small, 6-merous, hypanthium broad, P cucullate, disc with glandular projections, esp. alternating with K, A connective with a glandular tip, G 6, carpels plicate, loculi filled with secretions, 1(2) basal ovules/carpel; hypanthium accrescent in fruit; seed coat collapsed. - 1/1. S.W. Australia.

ELAEOCARPACEAE Candolle - Leaves simple; flowers pendant; C with 3 traces, large nectariferous disc/androgynophore, fringed, A many, centrifugal, basifixed, filaments short, anthers tubular-porose or with short slits, (connective prolonged), with lignified hairs, ovules ± hairy, outer integument 4-6 cells across, inner integument 6-12 cells across, style +, stigma ± punctate. - 12/625. Tropical, esp. Papuasia-Australia, not mainland Africa.


Leaf margin toothed [teeth with a single vein running into a congested ± deciduous apex]; stigma dry; exotegmen fibrous. - 39 families, 716 genera, 15935 species.

Achariaceae [Goupiaceae [Violaceae [Malesherbiaceae [Turneraceae + Passifloraceae]]] [Lacistemataceae + Salicaceae]]: sieve tubes with non-dispersive protein bodies; nectariferous tissue +, A = and opposite K, placentation parietal; endotegmen persistent.

ACHARIACEAE Harms - K and C not equal in number, and/or the perianth is not simply biseriate, disc 0; seed coat thick, testa vascularised, exotegmen massive. - 30/145. Pantropical.

[Goupiaceae [Violaceae [Malesherbiaceae [Turneraceae + Passifloraceae]]] [Lacistemataceae + Salicaceae]]: ?

GOUPIACEAE Miers - inflorescences umbellate; C induplicate-valvate, long, apical part inflexed, connective shortly prolonged, with long hairs, pollen with endexinal folds, placentation axile, several basal ovules/carpel, styles on outer shoulders of carpels; fruit a drupe; testa and tegmen ca 6 cells thick, testa with one layer [mesotestal] of sclereids, exotegmen poorly developed. - 1/2. N.E. South America.

VIOLACEAE Batsch - K quincuncial, A with abaxial nectary, G (3]; K persistent in fruit; exotesta subpalisade to tabular, ± thickened, (mesotesta sclerenchymatous), endotesta usu. crystalliferous. - 23/800. World-wide.

1. Fusipermoideae Hekking - C contorted, fleshy annular 5-lobed disc, filaments adnate to inner surface at indentations, A with minute fringed apical scales, thecae cordate/trapezoid, confluent apically?; capsule ca 2 mm long. - 1/3. Panama, Columbia, Peru.

2. Violoideae - C quincuncial, connectives prolonged, stigmatic head subcapitate, receptive area small. - 22/795. World-wide; woody taxa esp. in the lowland tropics.

Turneraceae + Malesherbiaceae + Passifloraceae: cyclopentenoid cyanogenic glucosides and/or cyclopentenyl fatty acids +; (plant with unpleasant smell); leaf teeth with vein proceeding to opaque and deciduous apex, (foliar glands +), colleters +; K + C together forming a tube, (corona or scales on C), A 5 [opposite K], well-developed stylodia +; funicular aril +, endotestal cells large, exotegmen palisade, endotegmen persistent.

Turneraceae + Malesherbiaceae: leaves spiral; micropyle bistomal; exotestal cells in lines; x = 7.

TURNERACEAE Candolle - 10/205. Tropical to warm temperate America and Africa (inc. Madagascar and Rodriguez I.).

MALESHERBIACEAE D. Don - K and C valvate; aril 0. - 1/24. Andean South America from Peru S., esp. N. Chile.

PASSIFLORACEAE Roussel - 17/575. Tropics to warm temperate, especially Africa and America.

SALICACEAE Mirbel - 55/1010. Pantropical, also temperate (but not Australia, New Zealand) to Arctic.

LACISTEMATACEAE Martius - P cup-like [1-6], A 1, the thecae ± separated and even stipitate, 1-2 apical ovules/carpel; aril ?0. - 2/14. Antilles, Mexico southwards, not in Chile.

Lophopyxidaceae + Putranjivaceae: stomata paracytic; flowers imperfect, 2 apical ovules/carpel; fruit 1-seeded.

LOPHOPYXIDACEAE H. Pfeiffer - Liane with leaf tendrils; secondary thickening with included phloem; K valvate, C very small, staminate flowers: A = and opposite K, cordate glands adnate to C; carpellate flowers: glands forming a lobed disc, G [5]; fruit a 5-winged samara. - 1/1. Malesia to the Solomon and Caroline Islands.

PUTRANJIVACEAE Endlicher - Glucosinolates +; plant dioecious; nucellus only ca 2 cells thick, disintegrating early, outer integument 3-8 cells and inner 6-14 cells across, archesporium 2-3-celled, endothelium +, stigmas flap-like; fruit a drupe; testa vascularised, exomesotesta sclereidal, tegmen 6-24 or more cells thick, exotegmen cells cuboidal. - 3/210. Tropical, esp. Africa and Malesia.

Ctenolophonaceae [Erythroxylaceae + Rhizophoraceae]: cristarque cells +; leaves opposite, stipules enclosing the terminal bud, interpetiolar; pedicels articulated; A 10, of two lengths, anthers with broad connective, 2 apical pendulous ovules/carpel; endosperm +.

CTENOLOPHONACEAE Exell & Mendonça - Leaves entire, stipules interpetiolar; C contorted, caducous, pollen 3-9 zonocolporate, G [2]; fruit with K persistent, swollen; seed single, persisting on columella; arillode ± hairy, exotestal cells palisade, the outer wall thickened. - 1/3. W. Africa, Malesia.

Erythroxylaceae + Rhizophoraceae: tropane [hygroline] and pyrrolidine alkaloids +; sieve tube plastids with protein crystalloids; mucilage cells +; stomata paracytic; leaves involute, colleters +; inflorescence cymose; K valvate, C conduplicate, enclosing A, A connate basally, inner integument ca 6 cells thick, endothelium +; fruit a septicidal capsule, K persistent; seeds also exotestal, arillate; endosperm starchy, embryo green.

ERYTHROXYLACEAE Kunth - 4/240. Pantropical, esp. American.

RHIZOPHORACEAE Persoon - Pits vestured; subepidermal laticifers in flower; C aristate, lobed, each enclosing one or more stamens, archesporial cells several. - 16/149. Pantropical.


1. Macariseae - Ca oxalate crystals solitary; stipules valvate; (seeds winged at micropylar end). - 7/94. Tropical America and Africa, aldo peninsula India and Sri Lanka.

Gynotrocheae + Rhizophoreae: stilt roots present; rootlets without root hairs; leaves bijugate; hypanthium +, ovary ± inferior.

2. Gynotrocheae - Fruit a berry; meso- and endotegmen persist. - 4/30: Crossostylis (10). Indo-Malesia, Madagascar.

3. Rhizophoreae - Stomata cyclocytic; abaxial hypodermis +; sclerenchymatous sheath of midrib at most weakly developed; leaves entire; endothelium 0; fruit indehiscent, 1-seeded; seed coat undifferentiated, vascularized, tegmen not persisting; seeds germinating on tree; cotyledonary node tri- or multilacunar. - 4/17 . Pantropical, but centred on the eastern Indian Ocean.

LINACEAE Perleb - C contorted, 2 apical ovules/carpel. - 10-12/300. World-wide.

1. Linoideae - C clawed. - 6/240. Worldwide, but esp. N. temperate and subtropical.

2. Hugonioideae - Stomata accessory cells usu. lignified, lobed beneath the guard cells; C yellow. - 4-6/61. Pantropical.

IRVINGIACEAE Exell & Mendonça - Cristarque cells widespread; stomata paracytic; leaves 2-ranked, revolute, margins entire, 2ndary veins strong, rather close and subparallel, tertiary veins also ± parallel, stipules large, intrapetiolar and encircling terminal bud; filaments folded in bud, disc massive, 1 pendulous ovule/carpel, placental obturator +; fruit indehiscent; testa thick, much sclerotised; cotyledons large, cordate. - 3/10. Africa; South East Asia to W. Malesia.

IXONANTHACEAE Miquel - Stomata paracytic; filaments folded in bud, pollen with supratectal spines, (1- Allantospermum)-2 apical ovules/carpel, placental obturator, endothelium +; fruit a septicidal capsule opening adaxially as well; endotegmen with sinuous anticlinal walls; cotyledons large. - 4-5/21. Pantropical.

HUMIRIACEAE A.-L. de Jussieu - Sieve element plastids with protein crystals and starch; filaments ± connate at least basally into tube, anther sacs separated, superposed, connective broad, prolonged; fruit a drupe, operculate, 1- or 2-seeded, surface sculpted; exotestal cells thick-walled, lignified, tegmen multiplicative. - 8/50. Tropical America, W. Africa.

PANDACEAE> Engler & Gilg - Cristarque cells +; plant dioecious: flowers small, nectary 0, 1 pendulous ovule/carpel; fruit a drupe. - 3/15. Tropics, Africa to New Guinea.

IRVINGIACEAE Exell & Mendonça - Stomata paracytic; stipules intrapetiolar, sheathing the stem; filaments plicate in bud; K thin, persistent in fruit, reflexed. - 3/10. Africa; South East Asia to W. Malesia.

Ochnaceae + Medusagynaceae + Quiinaceae [= Ochnaceae s.l.]: pits vestured; cristarque cells +; mucilage cellls/canals +; leaves with secondary and tertiary venation well developed; C contorted, nectary 0, ovules tenuinucellate.

OCHNACEAE de Candolle - Vessel/parenchyma pitting unilaterally compound [vessel side]; cortical vascular bundles +; anthers with pores; seeds winged; endotesta with small crystalliferous cells; endosperm slight. - 27/495. Tropical, esp. South America.

1. Luxembergioideae Endlicher - Androecium obliquely zygomorphic in bud, A developing adaxially only, filaments ± connate, anthers connate or not, pollen exine with small perforations. - 2/22. Venezuela and Brasil.

Ochnoideae + Sauvagesioideae: pollen with striate-rugulate exine.

2. Ochnoideae Burnett - Vessel/parenchyma pitting not unilaterally compound; one ovule/locule, integument single [connate, except sometimes at tip], 7-17 cells across; fruit indehiscent, usu. drupaceous; testa with vascular bundles, lacking layer of small crystalliferous cells, fibrous exotegmen 0; endosperm 0. - 9/290. Tropical, especially Brazil.

3. Sauvagesioideae Beilschmied - Exotesta with large cells, ± detached. - 16/82. Pantropical, only 2 spp. in Africa, most genera South American.

MEDUSAGYNACEAE Engler & Gilg - A many, G [16-25], attached to central axis, funicles long, stylodia on outer shoulders of carpels; fruit septicidal, carpels pulling away acropetally and opening adaxially, columella persistent. - 1/1. Seychelles.

QUIINACEAE Engler - Stomata anisocytic; fine venation paxillate. - 4/55. Tropical America.

[Clusiaceae + Bonnetiaceae] [Calophyllaceae [Hypericaceae + Podostemaceae]]: xanthones common; nodes 1:1; stomata paracytic; leaves with colleters, stipules 0; inflorescence cymose, C contorted, A many, nectary 0, micropyle bitegmic, stigma papillate; fruit a septicidal or -fragal capsule; exotegmen with anticlinal walls sinuous, low, lignified; endosperm at most slight, embryo ± fusiform.

Clusiaceae + Bonnetiaceae: ?

CLUSIACEAE Lindley, nom. cons.//GUTTIFERAE Jussieu, nom. cons. et nom. alt. - Leaves often flat, margins entire, linear canals +; plant dioecious; K and C decussate, style or styluli short; hypocotyl much enlarged, cotyledons minute. - 27/1050. Pan tropical.

BONNETIACEAE Nakai - 3/35. Malesia, Cuba, South America.

Calophyllaceae [Hypericaceae + Podostemaceae]: leaves with gland dots or lines.

CALOPHYLLACEAE J. Agardh - Androecium not obviously fasciculate, anthers with complex or simple glands; cotyledons moderate sized to huge. - 13/460. Throughout the tropics.

Hypericaceae + Podostemaceae: ovules tenuinucellate.

HYPERICACEAE Jussieu - 9/560. World-wide.

PODOSTEMACEAE Kunth - Annual (perennial) herbs of fast-flowing water, plant ± thalloid, stem root and leaf often not distinguishable, plant attached to substrate by haptera; primary root 0, other roots dorsiventrally flattened, shoots as endogenous buds from roots; SiO2 bodies +; pollen tricolpate, embryo sac monosporic, tetranucleate [Apinagia type], no polar nuclei or double fertilisation, nucellus plasmodial after fertilisation; capsule ribbed; exotesta thick-walled, often mucilaginous, endotegmen lignified; suspensor [micropylar] haustorium +; cotyledons large, plumule and radicle absent. - 48/270. Usually tropical, esp. America.

Podostemoideae + Weddellinoideae: G [2], with apical septum.

1. Podostemoideae - Flowers or groups of flowers enveloped by a spathella; microsporogenesis successive, pollen often in diads, outer integument develops first, nucellus plasmodial before fertilisation, style short, branches long; hypocotyl 0. - 43/260. Pantropical.

2. Weddellinoideae Engler - Capsule not ribbed; tegmen thick walled. - 1/1. N. South America.

3. Tristichoideae - Pollen pantoporate; hypocotyl 0. - 4/10. Tropical.

Centroplacaceae + Bhesa: inflorescence branched, pedicels articulated; A 5, opposite K, 2 ovules/carpel, styles widely diverging; exotegmen ribbon-shaped, thick-walled; embryo short.

CENTROPLACACEAE Doweld & Reveal - Stomata anisocytic; C 0, anthers basifixed, connective well developed, anthers with oblique-apical slits, pollen psilate. - 1/1. W. Africa.

Bhesa - Nodes 5:5; lamina with regularly ascending secondary veins and closely scalariform tertiary venation, petiole ± pulvinate apically, stipules almost encircling the stem; G [2]. - 1/5. Indo-Malesia.

Malpighiaceae + Elatinaceae: sieve tube plastids lacking starch and protein inclusions; leaves opposite, inflorescence cymose; nectary 0; fruit septifragal, K persistent; endosperm slight; x = 6.

MALPIGHIACEAE Jussieu - Hairs mesifixed; K with glands on abaxial surface, C clawed, crumpled in bud, one ovule/carpel. - 68/1250. Tropical and subtropical, especially American.

1. Malpighioideae - Tropical and subtropical, especially the Americas.

2. Byrsonimoideae - American Tropics.

ELATINACEAE Dumortier - 2/35. Worldwide, most tropical, not Arctic.

Peraceae + Rafflesiaceae + Euphorbiaceae: flowers small, imperfect; G [3], 1 apical pendulous epitropous ovule/carpel, nucellar cap + [unknown in Peraceae]; fruit a septicidal capsule/schizocarp, also splitting from the columella and loculicidally, mesocarp often separating from endocarp; seeds large, micropylar caruncle + (0); cotyledons longer and broader than radicle.

PERACEAE (Baillon) Klotzsch - Ovary septa membranaceous and without visible vascularisation; exotesta palisade, lignified, exotegmen tracheoidal. 4/135. Pantropical.

Rafflesiaceae + Euphorbiaceae: ?

RAFFLESIACEAE Dumortier - Stem or root parasites, rhizomes and roots 0; flowers large to massive; staminate flowers: A 12-40, adnate to central column, extrorse, anthers sessile, pores terminal, pollen inaperturate, atectate; carpellate flowers: G inferior, placentation laminar-parietal, ovules tenuinucellate, nucellar epidermis persists, stigma on outer margin or underside of disc-shaped structure; seed in two parts, that covered by the testa not enveloping the embryo, exotegmic cells with U thickenings, caruncle 0; endosperm slight, embryo undifferentiated. - 3/20. S. China, Assam, Bhutan, Thailand, W. Malesia, Australia.

EUPHORBIACEAE Jussieu - Exotegmen palisade. - 222/5970. Pantropical, also (warm) temperate. Four groups below, but these are only the well supported clades.

1. Peroideae Hasskarl - - 4/135. Pantropical.

1. Chelosioideae (Müller Argoviensis) K. Wurdack & Petra Hoffmann - Pollen echinate; testa with vascular bundles. - 2/7. Burma, Malesia.

Acalyphoideae [Crotonoideae + Euphorbioideae]: (phorbol esters [diterpenes] +); outer integument 6-10 cells or so across.

2. Acalyphoideae s. str. - Pantropical, some temperate.

Crotonoideae + Euphorbioideae: laticifers +.

3. Crotonoideae s. str. - Pollen inaperturate and with supractectal processes; tegmen with vascular bundles. - Pantropical, some temperate.

4. Euphorbioideae - Phorbol esters +; staminate flowers: P not enclosing A. - Pantropical, extending (mostly Euphorbia) into temperate regions.

Phyllanthaceae + Picrodendraceae: 2 apical ovules/carpel; fruit a septicidal capsule/schizocarp, also splitting from the columella and loculicidally, mesocarp often separating from endocarp, columella persistent.

PHYLLANTHACEAE Martynov - 56/1745. Pantropical, but esp. Malesia.

1. Phyllanthoideae Kostelevsky - Inflorescence fasciculate. - 38/ . Tropical to Temperate.

2. Antidesmatoideae Hurusawa - Plant tanniniferous; (pits vestured - Bridelieae); plant often dioecious; C often 0; fruit often indehiscent. - 21/ . Tropics and subtropics.

PICRODENDRACEAE Small - Stomatal subsidiary cells piggy back [on top of guard cells]; pollen spiny, nectary ?0. - 27/85. Many small genera; tropical, esp. New Guinea - Australia - New Caledonia (Caletieae) or America plus Africa - Madagascar (the rest).

Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae + Euphroniaceae]]: hairs simple [unicellular always?]; 2 ovules/carpel, micropyle bistomal; endosperm at most slight.

BALANOPACEAE Bentham & J. D. Hooker - Plant dioecious; staminate plant: inflorescence catkinate; P of small teeth, anthers much longer than filament; carpellate plant: inflorescence a fascicle, bracts spirally arranged and forming cupule; P 0, anthers much longer than filaments, ovules subbasal, apotropous, style branches long, once or twice bifid, stigmatic adaxially; fruit a drupe with 2-3 stones; testa vascularised, persistent, cell walls not much thickened; cotyledons cordate. - 1/9. S.W. Pacific, especially New Caledonia.

[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae + Euphroniaceae]: vestured pits +; mucilage cells +; stomata paracytic; leaf margins entire (surface glands or glandular hairs +); flowers obliquely monosymmetric; hypanthium +, K basally connate, quincuncial [?all], 2 outer members shorter, staminodes adaxial, fertile stamens abaxial, ± connate, connective well developed abaxially with endothecium continuous there, gynoecium with longitudinal furrows, unicellular unlignified hairs +, ovules epitropous, tenuinucellate, endothelium +, style +.

Trigoniaceae + Dichapetalaceae: (vessel elements with scalariform perforations); inflorescences cymose.

TRIGONIACEAE Endlicher - Branched sclereids +; hairs T-shaped, unicellular[?]; leaves with dense whitish hairs below; K connate, C contorted, adaxial-lateral = spurred standard, abaxial + abaxial-lateral = keel, A 5-13, filaments ± connate, fertile stamens 4-8, rest staminodial, pollen 3-5-porate, nectary glands [= staminodes?] at base of standard, inner integument to 12 cells across; capsule septicidal. - 5/28. Central and South America, Madagascar, W. Malesia.

DICHAPETALACEAE Baillon - Inflorescence epiphyllous, from the petiole; C bifid, drying black; fruit a flattened drupe, covered with short, erect hairs. - 3/165. Pantropical, few in Malesia.

Chrysobalanaceae + Euphroniaceae: hypanthium +.

CHRYSOBALANACEAE R. Brown - SiO2 bodies +; gynoecium borne on one side of the hypanthium, only one carpel functional, style ± gynobasic. - 17/460. Pantropical, especially American.

EUPHRONIACEAE Marcano-Berti - Lamina revolute; K unequal, C 3, A = and opposite K (-7), in two groups, adnate to C, filaments basally connate, staminodes 1, long, abaxial-lateral, and 4-5, small and dentate; capsule septicidal. - 1/1-2. The Guyana Shield, South America.

CARYOCARACEAE Voigt - Leaves trifoliolate; A many, filaments long, "glandular" at apex, G [4-20], styles separate; seeds reniform; hypocotyl very large, oily, spirally-twisted. - 2/21. Tropical America, esp. Amazonia.

CUCURBITALES + FAGALES [FABALES + ROSALES]: (N-fixing by root-dwelling associates [usu. the actinomycete Frankia]); seed exotestal; endosperm at most scanty, embryo large.

Cucurbitales + Fagales: G inferior; fruit 1-seeded, indehiscent.


Storied fusiform cambial initials +; perforation plates not or minimally bordered; rays wide, multiseriate; 2ndary veins palmate; K or P valvate, stomata on K/P raised, the two whorls rather similar in texture, micropyle bistomal, styles separate. - 8 families, 132 genera, 2320 species.

ANISOPHYLLEACEAE Ridley - Nodes 1:1; K epidermis with mucilaginous inner walls, C open, ± enclosing groups of A, lobed or laciniate, pollen heteropolar, dicolpate, psilate; seed coat initially thick, vascularised, becoming crushed; endosperm starchy, embryo fusiform, largely hypocotylar, cotyledons at most small. - 4/34. Pantropical.

[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Begoniaceae + Datiscaceae + Tetramelaceae]]: uniseriate rays 0; filaments shorter than anthers in bud, anthers basifixed, disc nectaries 0.

Corynocarpaceae + Coriariaceae: ellagic acid +; stomata paracytic; leaf margins entire; flowers small, petals thick, base broad, G superior, 1 apical ovule/carpel, vascular bundle extending into the outer integument; cotyledons very large.

CORYNOCARPACEAE Engler - Intrapetiolar stipules +; A = and opposite and basally adnate to C, incurved in bud, 5 fringed petaloid staminodes with a basal, adaxial nectary opposite K, pollen heteropolar, dicolpate, psilate, ektexine uniform, G [2], only one fertile. - 1/6. New Guinea to New Zealand, W. Pacific.

CORIARIACEAE Candolle - Nodes 1:1; leaves opposite; inflorescence racemose, bracteoles 0; C open, fleshy, G 5, at most only basally connate, styluli slender, stigmatic all around; fruit an achene, surrounded by accrescent C. - 1/5. Very disjunct: circum S. Pacific to China and Himalayas, Mediterranean.

Cucurbitaceae [Tetramelaceae [Datiscaceae + Begoniaceae]]: perennial herbs; cucurbitacins [triterpenes] +, myricetin, ellagic acid 0; young stem with separate bundles; leaves with teeth, medial vein ending in a pad of packed translucent cells, lateral also entering, stipules 0; flowers imperfect; a roof over the ovary [= stylodia marginal], stigmas large, elongated, bilobed; placentation parietal, many ovules/carpel.

CUCURBITACEAE Jussieu - Citrullin [non-protein amino acid - alpha-amino-delta-ureidopentanoic acid] +; tendrils +, bifid, lateral to the leaf; young stem with separate bicollateral vascular bundles; pericyclic sheath 0; cystoliths +; indumentum rough/prickly, walls calcified; C connate; staminate flowers: A extrorse, anthers monothecal, often much bent and coiled; carpellate flowers: placentae intrusive-parietal; fruit baccate, with a thick, hard skin; seeds flattened, pitted, testa multiplicative, exotesta complex; chalazal haustorium +. - 118/845. Especially tropical and subtropical.

1. Cucurbitoideae Kosteletzky - Often annuals; additional non-protein amino acids +; tendril branches alone coiled; staminate inflorescence + carpellate flower + bud + tendril making up axillary complex; plant monoecious; A inserted on hypanthium, pollen 40-200 µm long, style single; integument vascularised. - 101/745. Tropical to warm temperate.

2. Nhandiroboideae Kosteletzky - Plants perennial; tendril also coiled below the branches; plant dioecious; A on or near disc, pollen grains to 40µm long, striate, ovules pendulous, stylodia separate. - 18/70. Tropical.

Tetramelaceae [Datiscaceae + Begoniaceae]]: pollen spherical; fruit a capsule [dehiscing apically]; exotestal cells honeycomb, inner walls strongly thickened and lignified; cotyledons moderate in size.

TETRAMELACEAE Airy Shaw - Trees. - 2/2. Indo-Malesia.

Datiscaceae + Begoniaceae: ?

DATISCACEAE Berchtold & J. Presl - 1/2. W. North America, Crete to India.

BEGONIACEAE Berchtold & J. Presl - Herbs; stomata with accessory cells in two rings; pearl glands +; leaves 2-ranked, asymmetrical, stipules large, cauline-extrapetiolar; staminate flowers: A centrifugal, basifixed, connective enlarged, pollen colpate, carpellate flowers: borne towards the ends of the inflorescence branches. - 2/1401. Largely tropical.

1. Hillebrandia - Plant with round tubers; T 10 [K + C?], G [5], only partly inferior. - 1/1. Hawaii.

2. Begonia - Plant rhizomatous; staminate flowers: T in 2s; carpellate flowers: P 5, G [2-3], placentation axile, styles central; capsule dehiscing down sides, winged. - 1/1400. Largely tropical.

APODANTHACEAE Takhtajan - Parasites; plant monoecious or dioecious; P 2 + 4 + 4 or 3 + 6 + 6, nectary +, staminate flowers; gynostemium +, A synandrial, pollen sacs in rings, extrorse, pollen tricolpate, or apertures 0, psilate; carpellate flowers: G ± inferior, ovules tenuinucellate, style very short; fruit baccate; testa thin-walled, exo[?]tegmen massively lignified; endosperm +, embryo undifferentiated. - 3/23(+). New World from California and Florida southwards, Mediterranean and S. W. Asia, S. W. Australia and E. Africa.


Ectomycorrhizae common; dihydroflavonols; sieve tubes with non-dispersive P-protein bodies; bud scales +; leaf margins toothed, 2ndary veins proceeding straight to non-glandular teeth and higher-order veins convergent on those teeeth [urticoid]; plants monoecious, inflorescence spicate, flowers in compact cymose clusters [i.e. more than one flower/main inflorescence bract], very small; P +, (A opposite P), tectum ± spinulate, granular layer below tectum, nectary 0, 2 apical unitegmic ovules/carpel, poorly developed at pollination, integument 3-10 cells across, fertilisation delayed, style ± 0, stigmas ± decurrent; fruits dry; testa vascularised, not mechanical, exotesta often enlarged and persisting; cotyledons large. - 8 families, 55 genera, 1877 species.

NOTHOFAGACEAE Kuprianova - Peltate glandular hairs +; stipules peltate, enclosing colleters; fruit surrounded by cupule. - 1/35. New Guinea to South America.

Fagaceae + Myricaceae [Juglandaceae + Rhoipteleaceae] [Casuarinaceae + Ticodendraceae + Betulaceae]: anthers dorsifixed.

FAGACEAE Dumortier - Hairs often stellate/branched; sclereid nests with rhomboidal crystals in bark; cupule of valves [one more than fruit number], or continuous, scaly or spiny; pollen lacking granular layer below tectum; fruit surrounded by cupule of valves [one more than fruit number], ± spiny, individual nuts trigonous, endocarp hairy inside; seed pachychalazal. - 7/670. More or less worldwide.

1. Quercoideae - 6/640. N. temperate, at higher elevations in the tropics, not S. Africa or New Zealand, barely in Australia.

2. Fagoideae - Ellagic acid 0; inflorescence capitate; cotyledons folded. - 1/10. Temperate N. hemisphere.

[Myricaceae [Juglandaceae + Rhoipteleaceae]] [Casuarinaceae [Ticodendraceae + Betulaceae]]: myricetin +; pollen pororate, G [2]; fertilization chalazogamous.

[Myricaceae [Juglandaceae + Rhoipteleaceae]]: chains of crystal-containing cells in the wood; sieve tube P-protein bodies 0; peltate glandular hairs +; stipules 0; 1 flower/bract; ovule single [per flower], atropous, stigma lamellular/laciniate.

MYRICACEAE Kunth - Ovule basal, fertilisation porogamous. - 3/57. ± Cosmopolitan, including New Caledonia but not Australia.

Juglandaceae + Rhoipteleaceae: leaves odd-pinnate; P 4; endosperm 0; x = 16.

JUGLANDACEAE Perleb - Bracts expanded, bracteoles adnate to gynoecium; G unilocular above, ovule borne at apex of incomplete septum; pericarp intrusive; seeds large, pachychalazal, cotyledons much folded. - 7-10/50. North temperate, S. to Argentina and Malesia.

1. Engelhardioideae Iljinskaya - Leaves even-pinnate; bracts 3-lobed, bracteoles adnate to lower half of ovary; nuts with a layer of fibrous cells. - 3-4/14. Himalayas to Malesia, Mexico to Colombia.

2. Juglandoideae Manning - Vessel elements with simple perforation plates alone; staminate flowers: bracts 1-lobed, pollen usu. at least 26 µm [17-26 µm is the plesiomorphic condition] (with 4 or more pores); carpellate flowers: bracts unlobed, bracteoles usu. adnate to the ovary; fruit with sclereids in shell. - 3-6/35. Temperate N. hemisphere, only 1 sp. in Europe, Central America and Andes.

RHOIPTELEACEAE Handel-Mazzetti - Buds lacking scales; leaves 2-ranked, stipules +, asymmetrically caudate; 3 flowers/bract, central apparently perfect; pollen colporate, G [2], ovule bitegmic, campylotropous. - 1/1. China.

Casuarinaceae [Ticodendraceae + Betulaceae]: dihydroflavonols +[?]; stigmas elongate; fertilization chalazogamous.

CASUARINACEAE R. Brown - Rootlets clustered, of limited growth; nodes 1:1; stomata usu. tetracytic (hidden), transversely oriented; leaves 4-16-whorled, scale-like, margins entire, stipules 0; inflorescence capitate-spicate, 1 flower/bract; staminate flowers: P ["inner bracteoles"] 2, A 1, filaments incurved in bud, anthers ± longer than connective, pollen granules 0; carpellate flowers: bracteoles large, G naked, only abaxial fertile, ovules atropous, bitegmic, megasporocytes several, nucellar tracheids +; fruit a samara, freed as the much accrescent bracteoles separate. - 4/95. South East Asia and Malesia to the S.W. Pacific, esp. Australia.

Ticodendraceae + Betulaceae: sclereid nests with rhomboidal crystals in bark; mucilage cells +; leaves 2-ranked, doubly serrate.

TICODENDRACEAE Gomez-Laurito & L. D. Gomez P. - Hairs T-shaped, unicellular, not glandular; stipules encircling the stem; G with divided loculi, ovules hemitropous, integument 20-30 cells across. - 1/1. Central America.

BETULACEAE Gray - Flavones +; anthers longer than connective, pollen tube branched, horizontal transfer of rps11 gene. - 6/110. North Temperate, to Andes and Sumatra.

1. Betuloideae Arnott - No spirally-thickened vessel elements; peltate glandular hairs +; carpellate flowers: P 0; infructescence with woody or scaly bracts, separate fron small, ± flattened and samaroid nut. - 2/60. N. hemisphere, to South America; montane in tropics.

2. Coryloideae J. D. Hooker - Staminate flowers: P 0, A hairy; carpellate flowers: P +; infructescence with leafy bracteoles [from one or two orders of branching] remaining associated with the fruit; nuts large, not or little flattened. - 4/50. N. Temperate, South East Asia, Central America.

Fabales + Rosales: ?

FABALES Bromhead

Ellagic acid 0; styloids +; carpels free; embryo green. - 4 families, 754 genera, 20055 species.

QUILLAJACEAE D. Don - Nodes 1:3; hypanthium +, C clawed, antesepalous A on outer edge of the disc some way up the sepals, antepetalous A near base of G, G connate axially but not laterally, many ovules/carpel; fruit lobed, follicular, also opening abaxially; seeds winged; tegmen crushed; cotyledon investing radicle. - 1/3. Temperate South America.

Fabaceae [Surianaceae + Polygalaceae]: ?

FABACEAE Lindley//LEGUMINOSAE Jussieu - Non-protein amino acids and lectins [hemagglutinins] esp. in seeds +; floral developmental sequence K-G-C-outer whorl A-inner whorl A [G development much advanced], hypanthium +, C clawed, odd member of C adaxial, G 1, many ovules/carpel; fruit a follicle, also opening abaxially; exotesta palisade, linea lucida separating much thickened outer anticlinal walls from the rest, mesotesta of stellate cells, tegmen crushed; cotyledon investing radicle, endosperm ± starchy [Schleimendosperm]; rpl22 gene absent. - 730/19400. World-wide.

1a. Cercideae - Leaves apparently simple, bilobed or not. - 4-12/265. Pantropical (temperate).

1b. Duparquetia - Floral development acropetal ["normal"], K 4, petaloid, adaxial-median C external, A 4, opposite K, connate, porate, pollen asymmetrical, ectoaperture encircling the equator, with two endoapertures, G initiation not advanced. - 1/1. Tropical W. Africa.

1c. Detarieae - Resins with bicyclic terpenes; leaflets with crater-like glands on the abaxial surface, stipules deciduous; bracetoles decidous.

"Caesalpinioideae" + Mimosoideae + Faboideae: leaves bipinnate; vestured pits +.

2. "Caesalpinioideae" - 160/1930. Predominantly tropical, esp. Africa and America.

3. Mimosoideae - Sieve tube plastids also with fibres; extrafloral nectaries common; flowers of inflorescence develop together, bracteoles 0; flowers polysymmetrical, K connate, C connate, valvate, odd member abaxial, claws 0, A often connate; funicle long, thin, testa with vascular strand, pleurogram +, linea fissura [fine line delimiting pleurogram] common. - 82/3275. Esp. tropical and warm temperate, esp. Africa and America.

4. Faboideae - Isoflavonoids [pterocarpans and isoflavans]+; sieve tubes with spindle-shaped non-dispersive protein bodies; flower with sequence of development K-G-C-outer whorl A-inner whorl A, K initiation unidirectional, adaxial C external; hour-glass cells [below palisade exotesta] +, raphe shorter than the antiraphe, hilum with a hilar groove, tracheid bar [group of tracheids just below surface of hilum] +; embryo curved, with well-developed suspensor [?not the basal condition], radicle long, cotyledons not investing radicle, cotyledon areole +. - 476/13855. Esp. (warm) temperate, but world-wide.

Surianaceae + Polygalaceae: ?

SURIANACEAE Arnott - Carpels separate, styluli ± gynobasic. - 5/8 Mostly Australian, also Mexico, Suriana pantropical.

POLYGALACEAE Hoffmannsegg & Link - Flower monosymmetric; median abaxial C clawed, ± boat-shaped/keeled, A 8, median adaxial A 0, pollen polycolporate; K caducous, carpels connate; fruit a berry. - Ca 21/940 - four tribes below. World-wide, except the Arctic and New Zealand.

1. Xanthophylleae Chodat - Plants Al accumulators; glands at nodes, (conspicuous domatia on leaves), G [2], placentation parietal, 2 or more apotropous ovules/carpel, in two rows, outer integument 4-12 cells across, stigma small, bilobed (capitate); testa multiplicative; hypostase massive. - 1/95. Indo-Malesia.

The rest: A ± adnate to petals, variously connate, often monadelphous, anthers opening by short apical slits, 1 ovule/carpel, ovules epitropous.

2. Polygaleae Chodat - Flowers with two adaxial lateral K = wings, 2 abaxial lateral K, minute, two connate adaxial C = the standard, abaxial C = the keel, often fringed, 2 abaxial-lateral C minute, G [2], stigma complex, asymmetrical; fruit an often flattened capsule, drupe or samara. - Ca 13/830. World-wide, except Arctic and New Zealand.

3. Carpolobieae Eriksen - A (4) 5, G [3]. - 2/6. Tropical Africa.

4. Moutabeae Chodat

- Glands on leaves; seed with funicular aril. - 4/15. Tropical America, New Guinea to New Caledonia.

ROSALES Perleb  

Dihydroflavonols +; roots diarch [lateral roots 4-ranked]; prismatic crystals in ray cells; (sieve tubes with non-dispersive protein bodies; sieve tube plastids lacking starch); mucilage cells +; leaf margins with teeth; inflorescence cymose; hypanthium +, nectariferous, K valvate, C clawed, 1 apotropous ovule/carpel, micropyle endostomal, styles +, stigma dry; K and/or hypanthium persistent in fruit. - 9 families, 261 genera, 7725 species.

ROSACEAE Jussieu - A 10<, carpels separate. - 90/2520. World-wide, but esp. N. hemisphere.

1. Rosoideae - 2-pyrone-4,6dicarboxylic acid +; carpels many, 1 unitegmic, epitropous ovule/carpel; n = 7; plant with phragmidiaceous rusts. - Especially temperate (to Arctic) areas.

1A. Filipendula - Plant herbaceous; receptacle enlarged, 2 ovules/carpel. - 1/10. Eurasia.

Rosodeae T. Eriksson, Smedmark, & M. S. Kerr (= all other Rosoideae): ?

1B. Rubus - Prickly scrambling shrub; receptacle enlarged; fruit an aggregate of drupelets. - 1/± 250. ± Worldwide, esp. N. temperate.

1C. Colurieae Rydberg - 3/42. Temperate, inc. montane tropics, Chile.

[Rosa + Potentilleae] + Sanguisorbeae: ?

Rosa + Potentilleae: ?

1D. Rosa - Prickly arching shrub; 2 collateral ovules/carpel; hypanthium fleshy, urn-shaped. - 1/100-150. N. temperate.

1E. Potentilleae Sweet - Receptacle enlarged. - 11/800. N. temperate, esp. Europe, tropical mountains (S. temperate).

1F. Sanguisorbeae Candolle - Phragmidiaceous rusts 0. - 12/380. ± Worldwide, few Indo-Malesdia, tropical America.

Dryadoideae + Spiraeoideae: sugar alcohol sorbitol as transport carbohydrate, cyanogenic glycosides +.

2. Dryadoideae Juel - Association with N-fixing Frankia; ovules atropous; fruits achenes with hairy styles. - 4/19. W. North America, circumboreal (Dryas).

3. Spiraeoideae C. Agardh - Flavones +, ellagic acid 0; cuticle waxes as tubules or platelets; G <5, opposite C, 2< ovules/carpel, papillate funicular obturator +, stigma usu. wet; fruit a follicle.

3A. Lyonothamnus - Cyanogenic glycosides 0; leaves opposite, compound, stipules deciduous; G seminferior; ca 4 apical, epitropous ovules/carpel. - 1/1. California Islands, off S. California.

3B. Niellieae Maximowicz - Ovule single, apical; fruitlets hard, shiny. - 2/24. E. and W. North America.

3C. Amygdaleae Jussieu - Plant ectomycorrhizal; cork superficial; true tracheids 0; leaves conduplicate, nectaries on petiole or abaxial lamina; G 1, obturator from ovary wall; fruit a drupelet; n = 8. - 1/200. Temperate and tropical montane.

3D. Osmaronieae Rydberg - Cork superficial; pith chambered; stipules deciduous; ovules mid-position, obturator from ovary wall, styles lateral; fruit a drupe; n = 8. - 3/9. Central to East Asia, W. North America.

3E. Kerrieae Focke - Wart-like projections on lamina; ?obturator; fruit an aggregate, nut-like units. - 4/4. East Asia, W. North America, Alabama.

3F. Sorbarieae Rydberg - Leaves compound (simple: Adenostoma); ovules apical, epitropous; phragmidiaceous rusts 0. - 4/8. Central to East Asia, W. North America

3G. Spiraeeae Candolle - Vestured pits +; nodes 1:1 [?all]; stipules 0; >2 epitropous unitegmic ovules/carpel. - 8/106. N. temperate, to Columbia, (S. and) E. Africa, West Malesia.

Pyrodeae C. S. Campbell, R. C. Evans, D. R. Morgan, & T. A. Dickinson - Plant ectomycorrhizal; flavone C-glycosides +; cork superficial [?Gillenia]; rays often narrow; colleters + [probably elsewhere]; G ± connate, adnate to base of hypanthium, gynoecial ring primordium +, ovules basal, apotropous; exotesta ± thickened, often mucilaginous, mesotesta thick, sclerotic; Gymnosporangium rust common.

3H. Gillenia - Leaves compound. - 1/2. E. North America.

3I. Pyreae Baillon - N = 17; four copies of GBSSI [granule bound starch synthase I]. - 33/ca 1000.

Kageneckia + Lindleya - 4 pleurotropous ovules/carpel; Gymnosporangium rust 0. - 2/5. Mexico, Peru, Chile.

Vauquelinia - Tannin-containing cells pervasive; fruit septicidal, carpels opening adaxially (and partially abaxially as well). - 1/3. S.W. North America.

Pyrinae Dumortier - Stipules deciduous; G at leaf half inferior; hypanthium fleshy in fruit. - 30/1000. Mostly N. temperate to Arctic.

Barbeyaceae [Dirachmaceae + Rhamnaceae + Elaeagnaceae]: petiole bundle arcuate.

BARBEYACEAE Rendle - inflorescence fasciculate, bracts and bracteoles 0; flowers small, hypanthium and nectary 0, P uniseriate, anther with prolonged connective, G 1-2(-3), ± separate, ovule subapical, epitropous, style branches and stigmas long; fruit indehiscent, P accrescent. - 1/1. N.E. Africa, Arabia.

Dirachmaceae + Rhamnaceae + Elaeagnaceae: A = and opposite C/alternate with P, capsule septicidal; coat multiplicative, exotesta palisade, thick-walled; cotyledons large.

DIRACHMACEAE Hutchinson - Flower single, terminal, with epicalyx; C contorted, nectaries on base or on subbasal appendages, lacking stomata, A extrorse, anthers long, opening from apex; fruit beaked, segments opening adaxially, densely wooly inside, with columella. - 1/2. Socotra, Somalia.

RHAMNACEAE Jussieu - A enclosed by C. - 50/900. World-wide, especially tropics and warm temperate regions.

ELAEAGNACEAE Jussieu - Hairs lepidote or stellate; pits vestured; hypanthium long, C 0, G 1; hypanthium accrescent and fleshy in fruit; pericarp thin; endosperm with chalazal haustorium. - 3/45. North Temperate, warm tropical; Malesia and Australia.

Ulmaceae + Cannabaceae + Moraceae + Urticaceae: plant with ± watery exudate; hairs unicellular and multicellular-glandular; cambium ± storied; libriform fibers +; phloem stratified; sieve tubes with non-dispersive protein bodies; cystoliths [globose; usu. CaCO3] and epidermal and hair cell wall silicification and calcification; leaf with 2ndary veins proceeding straight to non-glandular teeth and higher-order veins convergent on those teeth [urticoid], at least one prominent prophyllar bud; flowers small, C 0, A = and opposite P, pollen porate, granular layer below tectum, nectary 0, G [2], abaxial only fertile, ovule apical, stigmas sessile, spreading, receptive area extending down adaxial surface and ± confluent; fruit a drupe; testa perforated[?]; x = 14, centromeres both median and subterminal.

ULMACEAE Mirbel - Seeds flattened. - 6/35. Mostly N. temperate, esp. Asian, but scattered elsewhere except Australia and the Pacific.

Cannabaceae + Moraceae + Urticaceae: C-glycoflavones also +; unicellular hairs usu. micropapillate; 2ndary veins palmate; stipules cauline-intrapetiolar; flowers imperfect; embryo curved.

CANNABACEAE Martynov - 11/170. Worldwide, but not Arctic.

Moraceae + Urticaceae: latex system +; stamens inflexed in bud.

MORACEAE Link - Laticifers throughout plant, latex milky; inflorescence compact. - 38/1100. Mostly tropical to warm temperate.

URTICACEAE Jussieu - Laticifers only in bark; lignification of vascular tissue delayed; G apparently 1, ovule basal, ± atropous. - 54/2625. World-wide, but mainly tropical.


[GERANIALES + MYRTALES]: ellagic acid +; K persistent in fruit[!]


Ellagic acid +; leaf margins gland-toothed; inflorescence cymose; nectary outside A, A obdiplostemonous, G opposite C, micropyle bitegmic; K persistent in fruit; seed testal. - 5 families, 17 genera, 836 species.

GERANIACEAE Jussieu - 7/805. Temperate and warm temperate.

1. Hypseocharis - Stipules 0; A 15, style filform, stigma capitate; fruit a loculicidal capsule, filaments also persistent; seeds with scanty endosperm, cotyledons spiral. - 1/1-3. S.W. Andean South America.

2. The rest - K with nectariferous spurs, aristate; 1-2 apical ovules/carpel, style short, stout, stigma lobed; fruit with upper part of ovary elongating [the "stylar" beak], separating into mericarps which curl upwards and separate from columella, whether or not seeds disperse separately; embryo curved, cotyledons longitudinally folded. - 6/805. Temperate and warm temperate, esp. southern Africa.

Melianthaceae + Francoaceae: at least traces of inulin +; leaves conduplicate, spiral, insertion rather broad; inflorescence terminal, racemose, sterile bract(s) at apex, bracteoles 0; G lobed, placentae intrusive, ovules in two rows, style long; endosperm copious, embryo short.

MELIANTHACEAE Berchtold & J. Presl - 3/11. Africa.

  • Melianthaceae are at most small trees that may be recognised by their serrate leaves or leaflets, and large, toothed stipules. They have terminal racemes of large, more or less monosymmetric flowers with all the parts (apart from the ovary and sometimes calyx) free; the nectary is outside the stamens.

FRANCOACEAE A. Jussieu - 2/2. Chile.

Vivianiaceae + Ledocarpaceae: leaves opposite, simple (margins entire), stipules 0, but a line across the stem; inflorescence terminal; K aristate, C contorted, pollen grains polypantoporate, micropyle endostomal, funicle with hairs, style short, stigmatic lobes long; fruit a loculicidal capsule; seed coat more or less tanniniferous; endosperm walls thick, pitted, embryo curved.

VIVIANACEAE Klotzsch - Nodes 1:1. - 1 or 4/6. Chile, S. Brasil.

LEDOCARPACEAE Meyen - 3/12. W. South America, especially the Andes.

MYRTALES Reichenbach  

Flaky bark; flavonols only, myricetin, methylated ellagic acid +; cork pericyclic; pits vestured; libriform septate fibers +; secondary phloem stratified in young twigs; internal phloem +; nodes 1:1; leaves opposite, (small stipules +), colleters +; hypanthium +, nectariferous, K valvate, A incurved in bud, pollen with pseudocolpi, many ovules/carpel, micropyle bistomal and zig-zag, style single, long, minor stylar bundles +; K persistent in fruit; endotesta crystalliferous; endosperm slight. - 11 families, 380 genera, 11027 species.

COMBRETACEAE R. Brown - Hairs with a basal internal compartment; ovary unilocular, ovules apical; fruits indehiscent, ± flattened and/or winged; seed large. - 14/500. Largely tropical.

Onagraceae + Lythraceae: tannins often not abundant, soluble oxalate accumulating; vessels in groups in the wood; fibers with at most minutely bordered pits; petiole bundle arcuate; (flowers vertically monosymmetric), C clawed, (pollen at anthesis with starch), several megasporocytes; K persistent in fruit; fibrous exotegmen +; starch grains in nucellus; n = 7.

ONAGRACEAE Jussieu - Raphides +; A straight, anthers locellate, pollen with viscin threads, exine paracrystalline, beaded, embryo sac 4-nucleate, stigma capitate; fruit a capsule opening down the sides; endosperm nuclear, diploid. - 18/650. World-wide, esp. America.

1. Jussiaeaoideae - Hypanthium 0, pollen in tetrads (monads), nectary on G, G with central vascular bundles, style short. - 1/85. World-wide, esp. America.

2. Onagroideae - Flowers 4-merous; ypanthium long, transseptal vasular bundles +, no minor stylar bundles; K not persistent in fruit. - 17/565. World-wide, esp. S.W. North America.

LYTHRACEAE Jaume Saint-Hilaire - C crumpled in bud; testa many layered. - 31/620. Tropical, but some temperate.

[Vochysiaceae + Myrtaceae] [Melastomataceae + Penaeaceae]: ?

Vochysiaceae + Myrtaceae: hairs simple, 1-2-celled; K and C imbricate, pollen syncolporate; style depressed in apex of gynoecium.

VOCHYSIACEAE A. Saint-Hilaire - Flowers obliquely monosymmetric, K with a nectar spur, A 1. - 7/190. Lowland tropical America, W. Africa.

MYRTACEAE Jussieu - Aromatic compounds [terpenes, etc.] +; wood fibers with distinctly bordered pits; leaves with gland dots. - 131/4625. Tropical and (warm) temperate.

1. Psiloxyloideae (Croizat) R. Schmid - Leaves spiral; plant dioecious; C clawed, staminate flowers: A erect in bud, with separate traces, usu. 4 thecae opening separately, pistillode +; carpellate flowers: staminodia +, G [3], embryo sac bisporic, 8-nucleate; endotesta cells periclinally elongated. - 2/4. Southern Africa, Mascarenes.

1A. Heteropyxideae Harvey - A = and opposite K[?] (+ 2 or 3 opposite C), G fully superior [base narrow]. - 1/3. S.E. Africa.

1B. Psiloxyleae (Croizat) A. J. Scott - Style 0; fruit a berry. - 1/1. Mascarenes.

2. Myrtoideae Sweet - Stamens many, with terpenoid-producing glands. - 129/4620. Tropical, also temperate, esp in Australia.

Melastomataceae + Penaeaceae: (plants Al accumulators); (nodes swollen); branched or unbranched sclereids +/0 within same family; inflorescence cymose; connective abaxially much expanded [least in Rhynchocalycaceae], disc 0.

MELASTOMATACEAE Jussieu - Included phloem +; C contorted, anthers with branched vascular trace, connective expansion below anther well developed, with various appendages; radicle bent. - 194/5000. Pantropical (warm temperate).

?name of clade - Lamina with veins diverging at the base, tertiaries scalariform; A held on one side of the flower, connective basally thickened and with a ± prominent appendage. - 182/4570. Largely tropical and subtropical, esp. South America.

1. Pternandra - Hairs uniseriate; endothecium in inner wall of inner sporangium only, placentation parietal; capsule fleshy. - 1/15. Southeast Asia.

2. Melastomatoideae - Included phloem 0; leaf veins lacking fibrous sheath; endothecium ephemeral.

2A. Astronieae - Hairs as peltate scales; placentation basal to basal-axile. - 4/150. Indomalesia and Pacific.

2B. REST - 177/4305. Largely tropical and subtropical, esp. South America.

Olisbeoideae - Sclereids +[?]; anthers with oil-secreting gland. - 6/435. Tropical.

PENAEACEAE Guillemin - Stems square, nodes swollen, stipules minute; A = and opposite C, endothecium ephemeral; fruit a capsule; exotestal cells periclinally elongated, endotegmen not fibrous.

Crypteronioideae van Beusekom - 3/10. South East Asia, Malesia, Sri Lanka.

Penaeaoideae - Style stout. - 9/25. Southern and eastern Africa, Costa Rica to Peru.



?Nodes; leaf margins toothed, stipules cauline; hypanthium +, with nectary [disc], G stipitate, ovules apotropous, micropyle bistomal [zig-zag], both integuments 4 or more cells across, styles free, compitum present only near stigma, stigma wet; seed testal, walls of outer layers much thickened. 7 families, 12 genera, 66 species.

Staphyleaceae [Guamatelaceae [Stachyuraceae + Crossosomataceae]]: leaves or leaflets involute; stigma ± expanded.

STAPHYLEACEAE Martynov - Leaves opposite, odd-pinnate, with small "glands" at the articulations; A = and opposite K. - 2/45. North temperate, tropics in America (to Bolivia) and Malesia.

Guamatelaceae [Stachyuraceae + Crossosomataceae]: funicular aril +.

GUAMATELACEAE S. Oh & D. Potter - Leaves opposite, venation palmate; pollen membranes protruding through apertures; endosperm scanty. - 1/1. Mexico to Honduras.

Stachyuraceae + Crossosomataceae: crystals/druses absent from flowers, anthers X-shaped; whole testa sclerotic.

STACHYURACEAE J. Agardh - Pith well developed; stomata paracytic; inflorescence long-racemose; flowers 4-merous, hypanthium 0, nectary at base of G, G [4], placentation axile, style +; fruit a berry. 1/5-10. South East Asia.

CROSSOSOMATACEAE Engler - 4/12. W. North America.

Aphloiaceae [Geissolomataceae + Strasburgeriaceae]: conspicuous protrusions from pollen apertures.

APHLOIACEAE Takhtajan - Cork pericyclic; stomata anisocytic; inflorescence fasciculate; P ca 7, ?spiral, A many, pollen striate, G 1, micropyle endostomal, stigma sessile, annular-peltate; fruit a berry. - 1/1 or more. E. Africa, Madagascar, the Mascarenes and Seychelles.

Geissolomataceae + Strasburgeriaceae: hairs single-celled and T-shaped; pollen ± psilate; G not stipitate, with abaxial [dorsal] ribs, two collateral ovules/carpel, stigma punctiform; fruit a capsule; hilum on seed large.

GEISSOLOMATACEAE Endlicher - Nodes 1:1; flowers axillary, 4-merous, with 3-4 pairs of basal bracts, P 4, petaloid, basally connate, A adnate to base of P. - 1/1. Cape Province, South Africa.

STRASBURGERIACEAE Solereder - Cells with thickened mucilaginous inner tangential walls [in flowers, at least]; acicular crystals +; leaves spiral, margins gland-toothed; flowers large; K spiral, C clawed, filaments flattened, anthers >3mm long, ovules sessile, epitropous, style long; style persistent in fruit; x = 25. - 2/2. New Caledonia, New Zealand.


The position of the character of ovule number on the tree is unclear. However, taxa with one or two ovules/carpel are common in the rosid II clade, and the ovules are usually apical.


C18 acetylenic acids [tariric, petroselenic], anthraquinones +. - 1 families, 2 genera, 46 species.

PICRAMNIACEAE Fernando & Quinn - Leaves odd-pinnate, stipules 0; A = and opposite C; 2 ovules/carpel. - 2/46. Neotropical.

1. Picramnioideae Engler - Ovules apical; fruit a berry. - 1/41. S.E. U.S.A., Central and South America, Caribbean.

2. Alvaradooideae Liebm. - Staminodes opposite K, only 1 G fertile, ovules basal, apotropous; fruit a samaroid capsule. - 1/5. Florida, Central America, Bahama, esp. the Greater Antilles, Bolivia to Argentina.

SAPINDALES [HUERTEALES [MALVALES + BRASSICALES]]: ?flavonols +; vessel elements with simple perforation plates; petiole bundle(s) annular; inner integument thicker than outer, style +; endosperm scanty.

SAPINDALES Dumortier  

Interesting secondary compounds, ethereal oils, myricetin +; silicified wood or wood with SiO2 grains; leaves spiral, odd-pinnately compound, conduplicate; intrastaminal nectariferous disc well developed; seed coat?. - 9 families, 460 genera, 5670 species.

BIEBERSTEINIACEAE Endlicher - 1 apical pendulous ovule/carpel, embryo sac tetrasporic, 16-nuclear, styles separate, impressed, apically connate, stigma capitate; fruit a schizocarp, columella persisting. - 1/5. Greece to Central Asia.

NITRARIACEAE Berchtold & J. Presl - 3/16. Usu. ± arid regions in N. hemisphere, Australia.

Kirkiaceae + Anacardiaceae + Burseraceae: 2 ovules/carpel; fruits with 1 seed/carpel.

KIRKIACEAE Takhtajan - Inflorescence dichasial, flowers small, 4-merous, staminate flowers: stamens = and opposite sepals, pollen syncolpate; carpellate flowers: G [8 (always?)], 1 ovule/loculus, tip of axis in ovary swollen, glandular, stigma with post-genital fusion, stigmas covered with large uniseriate hairs; fruit a schizocarp, mericarps pendulous from columella. - 2/6. Tropical and S. Africa.

Anacardiaceae + Burseraceae: biflavones +; vertical intercellular secretory canals in phloem, this surrounded by a light-coloured, sinuous, sclerenchymatous band; flowers rather small, K often connate, C little longer than K, A obdiplostemonous; fruit a drupe, endocarp cells lignified, not oriented.

ANACARDIACEAE R. Brown - 1 ovule/carpel. - 70/985. Tropical, also temperate.

1. Spondoideae Arnott - Biflavonoids; also alkylcathechols and alkylresorcinols 0; G [(1-[pseudomonomerous?])4-5(-12)] and as many locules, ovule pendulous; exocarp thick. - 10/115. Tropical.

2. Anacardioideae Link - (Biflavonoids, 5-deoxyflavonoids, also alkylcathechols and alkylresorcinols [phenols with unsaturated side chains - allergenic] +); leaves simple (compound); antesepalous carpel alone fertile; exocarp thin, epidermis lignified, endocarp with up to three layers of palisade lignified sclereids, internal to these a crystalliferous layer [= stratified]. - 60/485. Largely tropical, also temperate.

BURSERACEAE Kunth - 18/550. Tropical, but esp. America and N.E. Africa.

1. Beiselieae Thulin, Beier & Razafimandimbison - Leaf ?bases persistent, with a spine; G [9-12], ovules superposed; fruit septifragal, pericarp splitting septifragally separately, columella with deep flanges, mericarps apically winged. - 1/1. Mexico.

2. Bursereae DC, Garugeae Marchand, Protieae Marchand - Stigmatic head formed by postgenital fusion; cotyledons folded, palmately lobed. - 17/550. Tropical, but esp. America and N.E. Africa.

SAPINDACEAE Jussieu - Cyclopropane amino acids +; C with various adaxial appendages/folds, disc outside A, A 8 or fewer, G [3] [/level], ovules sessile; testa vascularized; embryo curved, the radicle in a pocket of the testa. - 135/1580. ± World-wide.

1. Xanthoceroideae Thorne & Reveal - Leaves deciduous; disc with golden, horn-like glands, 6-8 ovules/carpel. - 1/1. N. China.

Hippocastanoideae + Dodonaeoideae + Sapindoideae: flowers often strongly obliquely or vertically [Aesculus] monosymmetric, 2 ovules/carpel.

2. Hippocastanoideae Burnett - Leaves opposite, palmate. - 5/130. N. temperate, some tropical, usu. mountains.

3. Dodonaeoideae Burnett - Pantropical-warm temperate, esp. Australia/Southeast Asia.

4. Sapindoideae Burnett - 60/1 (2) ovules/carpel. - Pantropical. Paullineae (8 genera) contain one third of the species in the family.

Rutaceae + Meliaceae + Simaroubaceae: alkaloids, pentanortriterpenes +; inflorescence branches cymose; disc between A and G.

Rutaceae + Meliaceae: tetranortriterpenes, flavones +; stigma capitate.

RUTACEAE Jussieu - Leaves punctate. - 161/1815. Largely tropical.

MELIACEAE Jussieu - Filaments connate, stigmatic head expanded. - 52/621. Pantropical, but largely Old World; plants of the lowlands.

1. Melioideae Arnott - Seeds with arillode or sarcotesta. - 37/571. Pantropical, but largely Old World.

2. Swietenioideae Arnott - Buds perulate; 3-many ovules/carpel; fruit a septifragal capsule, valves falling off, columella persisting. - 15/50. Pantropical, but largely Old World.

SIMAROUBACEAE Candolle - Quassinoids +; carpels united by styles alone, 1 ovule/carpel; fruitlets separate, drupes. - 19/95. Largely tropical; a few (e.g. Ailanthus) temperate.

Huerteales [Brassicales + Malvales]: ?


Vessel elements with scalariform perforations; flowers small, A = and opposite K, ovules 1-2/carpel; endosperm copious, embryo at most medium. - 3 families, 5 genera, 21 species.

TAPISCIACEAE Takhtajan - Leaves compound, articulated, with small "glands" at the points of articulation; pollen colpate. - 2/5. China, West Indies and N. South America.

GERRARDINACEAE Alford - Hypanthium +, A opposite C, G unilocular, placentae apical, embryo minute. - 1/2. Eastern Africa.

DIPENTODONTACEAE Merrill - Inflorescence umbellate; K and C similar. - 1/1. S. China and adjacent Burma.

Brassicales + Malvales: ?


Idioblastic and stomatal myrosin cells +, glucosinolates from phenyalanine and/or tyrosine; endoplasmic reticulum with dilated cisternae; myricetin and other methylated flavonols, tannins 0; leaves spiral; inflorescence racemose; (petals clawed), ovules in one or two rows; seed coat? - 17 families, 398 genera, 4450 species.

Akaniaceae + Tropaeolaceae: vessel elements with scalariform perforations; bracteoles 0; flowers quite large, obliquely monosymmetric in bud, K/C tube +, C clawed, A 8, with short connective prolongation, placentation apical-axile, 1-2 epitropous ovules/carpel, style long; testa vascularised.

AKANIACEAE Stapf - Leaves pinnate; flowers obliquely asymmetric when open. - 2/2. S.W. China, adjacent Vietnam, Formosa, E. Australia.

TROPAEOLACEAE Berchtold & J. Presl - Rather fleshy vines; venation palmate; flowers monosymmetrical, with adaxial calycine spur, ovules tenuinucellate; fruit a schizocarp. - 1/95. New World.

[Moringaceae + Caricaceae] [Setchellanthaceae [Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae [Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]]]]]]: ?

Moringaceae + Caricaceae: woody, stems stout; endoplasmic reticulum-dependent vacuoles; venation palmate, colleters +, stipules as glands; inflorescences thyrses; flowers whitish, ovary longitudinally sulcate, replum stands opposite the ventral bundles +[?], placentation parietal, many ovules/carpel, outer integument 5-6 cells across, micropyle bistomal, style hollow; testa multiplicative, mesotesta ± lignified.

MORINGACEAE Martynov - Schizogenous gum canals +; leaves odd-pinnate, 1- or 3-compound, with glands at the articulations and at the base; flowers obliquely monosymmetric, median C abaxial, A = and opposite C, declinate, unithecal, staminodes opposite K; fruit a long, slender capsule; seeds 3-angled. - 1/12. India to Africa, Madagascar.

CARICACEAE Burnett - Articulated laticifers +, anastomosing; leaves palmately-veined or strongly lobed (palmate), glands on adaxial surface at base; plants di(mon)oecious; staminate flower: C connate, A adnate to C, nectary on pistillode; carpellate flowers: C often free, nectary 0, G [5], styles ± separate; fruit a berry; sarcotesta +, mucilaginous, mesotesta tanniniferous, with lignified ribs, endotesta crystalliferous. - 4(-6)/34. Mostly tropical America; Africa (Cylicomorpha only).

Setchellanthaceae [Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae [Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]]]]]: nodes 1:1; extended 3' terminus of rbcL gene.

SETCHELLANTHACEAE Iltis - Hairs T-shaped; flowers 6-merous, K opening irregularly; capsule septifragal, columella persistent. - 1/1. Mexico.

Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae [Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]]]]: root hairs in vertical files.

LIMNANTHACEAE R. Brown - 1 basal tenuinucellate unitegmic ovule/carpel, embryo sac tetrasporic, 4- or 6-nucleate, style gynobasic; fruit a schizocarp, mericarps muriculate; seed coat pachychalazal, thick, with vascular bundles, otherwise undistinguished; cotyledons cordate. - 1(2)/8. Temperate North America.

[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae [Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]]]: ovules campylotropous, style/styles short to absent; seeds exotegmic, exotegmen fibrous; embryo strongly curved.

Koeberliniaceae [Bataceae + Salvadoraceae]: idioblastic myrosin cells 0; flowers 4-merous, pollen 3-colporoidate, disc 0, G [2]; exotestal cells well developed; n = 11.

KOEBERLINIACEAE Engler - Plant thorny, leaves reduced; ovule tenuinucellate, nucellar epidermal cells radially enlarged, outer integument 2 cells across, style long. - 1/1. C. and S.W. North America, Bolivia(!).

Bataceae + Salvadoraceae: rays wide, multiseriate; nodes two trace; stomata paracytic; leaves opposite, with 2ndary veins ascending from at or near base; pollen psilate, G (2), 2 basal ovules/carpel; tegmen not fibrous; endosperm 0, embryo ± straight, color?

BATACEAE Perleb - Leaves fleshy; inflorescence more or less strobilate; flowers imperfect; carpellate flowers: P 0, G 4-locular. - 1/2. N. Australia and S. New Guinea, tropical America, and the Galapagos.

SALVADORACEAE Lindley - 3/11. Africa (inc. Madagascar) to South East Asia and West Malesia, often in drier regions.

Emblingiaceae [Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]]: glucosinolates also from valine/isoleucine and/or leucine, indole glucosinolates from tryptophan; cisternae of endoplasmic reticulum dilated and vacuole-like; inflorescence terminal, racemose, bracteoles 0; floral development open, C clawed, disc/nectary outside A, ovules ± campylotropous; exotegmen fibrous, endotesta crystalliferous; 3' rbcL extension.

EMBLINGIACEAE Airy Shaw - Flowers axillary, monosymmetric, resupinate, K connate, lobed, deeply divided adaxially, C 2, ?not clawed, abaxial, connate by epidermis, slipper-shaped, nectary abaxial, androgynophore curved abaxially, A 8, median members absent, 4 abaxial fertile, 4 adaxial staminodial, forming a torus. - 1/1. W. Australia.

Pentadiplandraceae [Gyrostemonaceae + Resedaceae] Tovariaceae [Cleomaceae [Capparaceae + Brassicaceae]]: ?.

PENTADIPLANDRACEAE Hutchinson & Dalziel - K 5, valvate, C 5, connivent at enlarged, concave base, limb flat; carpellate flowers: style long, stigma shortly lobed; fruit a berry; 1 seed/loculus, coat with layer of white, wooly, elongated cells towards outside. - 1/?1. Tropical W. Africa.

Gyrostemonaceae + Resedaceae: hairs unicellular; styluli +; seeds arillate.

GYROSTEMONACEAE Endlicher - P uniseriate, connate, pollen colpate, G borne around the central axis in whorl(s). - 5/18+. Australia.

RESEDACEAE Berchtold & J. Presl - C ligulate at junction of claw and limb, limb fringed. - 6/75. N. warm temperate and subtropical, esp. Mediterranean-Middle East-North African, also Africa.

TOVARIACEAE Pax - 1/2. Tropical America.

Capparaceae [Cleomaceae + Brassicaceae]: sinapine [methyl glucosinolate], erucic acid [fatty acid] +; cisternae of endoplasmic reticulum organelle-like, etc.; lateral wall pits of vessels vestured; eglandular hairs simple, unicellular [?level]; flowers 4-merous, A from 4 primordia, centrifugal, longer than the petals, gynophore +, placentation parietal; K deciduous; seeds with invaginated coat.

CAPPARACEAE Pax - Fruit indehiscent. - 16/480. Largely tropical.

Cleomaceae + Brassicaceae: herbaceous annuals; inflorescence ± corymbose; C 4, clawed, A 6, G [2]; fruit septicidal, persistent, woody placenta +.

CLEOMACEAE Horaninow - Anthers linear, coiled at dehiscence. - 10/300. Tropical and warm temperate, esp. America.

BRASSICACEAE Burnett - Methyl glucosinolates 0; roots lacking mycorrhizae; the two outer A shorter than the four inner, about as long as petals, commissural septum +, stigma commissural; seed folded, but no invagination of the coat; chalazal endosperm cyst +; duplication of PHYB -> PHYD gene. - 338/3710. World-wide, esp. N. temperate (but less E. North America).

Aethionemeae Al-Shehbaz, Beilstein & E. A. Kellogg - Plant glabrous; fruit angustiseptate. - 1-2/70. The Mediterranean and Europe to Afghanistan.

The Rest - Genome duplication. - 335/3640. World-wide, esp. N. temperate (but less E. North America, and even more so humid lowland tropics).

Malvales + Brassicales: ?

MALVALES Dumortier  

(Cyclopropenoid fatty acids +), flavones, myricetin +; mucilage cells +; stipules +; C contorted, nectariferous disc 0, few ovules/carpel, style long; exotegmen much thickened and lignified, palisade. - 10 families, 338 genera, 6005 species.

NEURADACEAE Link - Pollen bipolar, with 3 or 4 pores at each end, not all carpels fertile, styles separate. - 3/10. Africa to India, dry or desert areas.

Thymelaeaceae, Sphaerosepalaceae, Bixaceae, [Cistaceae + Sarcolaenaceae + Dipterocarpaceae], [Cytinaceae + Muntingiaceae], Malvaceae: pits vestured; phloem stratified, phloem rays wedge-shaped.

THYMELAEACEAE Jussieu - Pericyclic fibers 0; crystal sand +; epidermal cells (massively) mucilaginous; stomata cyclocytic. - 46-50/891. World-wide, esp. trop. Africa and Australia.

1. Tepuianthus - 1/7. Guayana highlands, N.E. South America.

Octolepidoideae + Thymelaeoideae: C 0, P [= K?] tubular, with apical appendages, pollen oligo- to polyporate, minutely spinulose, style single, stigma ± capitate, dry; endotegmen with stripes on the inner surface.

2. Octolepidoideae Gilg - Secretory cavities + [leaves punctate]; glandular scales 4-40, anthers usu. recurved and hippocrepiform, connective well developed, G (2-)3-5(-8); seed arillate, with swollen funicle, or angled at the raphe; chalazal fold on ventral side only [distribution elsewhere in family?], nucellar tracheids +. - 8/49. Tropical Africa and Madagascar Octolepis, Malesia to Australia (Queensland), New Caledonia, Fiji.

3. Thymelaeoideae Burnett - Phorbol ester diterpenes, chelidonic acid +; internal phloem +; vascular bundles bicollateral; inflorescence often capitate; hypanthium long, petaloid appendages to 2 x K or 0, A 2-5 or 10, pollen crotonoid, with supratectal processes, G 2; fruit a drupe or achene. - 37/690. World-wide, esp. trop. Africa and Australia.

Sphaerosepalaceae + Bixaceae [Cistaceae + Sarcolaenaceae + Dipterocarpaceae] + Muntingiaceae + Malvaceae: hairs often stellate; stipules often well developed; A many, developing centrifugally, from 5 or 10 (15) bundles, when 5 often opposite the petals, several [³6] ovules/carpel, micropyle bistomal.

SPHAEROSEPALACEAE Bullock - 2/18. Madagascar.

BIXACEAE Kunth - Plant with secretory canals; resin-filled cells outside veins; hairs glandular, not tufted or stellate; leaf teeth with a single vein proceeding to opaque deciduous apex; inflorescence terminal, flowers large; K imbricate, anthers porate, many ovules/carpel, funicles long, micropyle zig-zag, stigma at most slightly lobed; exotegmen curved inwards in chalazal region, hypostase plug with core and annulus, outer hypostase forming the core; embryo long, cotyledons spatulate, thin, curved or folded, radicle short, stout. - 4/21. Pantropical.

Cistaceae + Sarcolaenaceae + Dipterocarpaceae: ectomycorrhizae +; ellagic acid +; plant with secretory canals; K imbricate, two outer members often different from the rest, filaments not articulated, ovules both anatropous and atropous; exotegmen curved inwards in chalazal region, hypostase plug with core and annulus; endosperm starchy.

CISTACEAE Jussieu - Stigma with multicellular multiseriate papillae. - 8/175. More or less worldwide, often temperate or warm temperate, esp. Mediterranean region.

Sarcolaenaceae + Dipterocarpaceae: petiole anatomy complex; stipules usu. well developed.

SARCOLAENACEAE Caruel - Involucre of varying morphology subtending flowers; disc extrastaminal, pollen in tetrads. - 8/60. Madagascar, mostly E. and C., fossil pollen in South Africa.

DIPTEROCARPACEAE Blume - K enlarging equally, thinnish; testa vascularized. - 17/680. Tropical, overwhelmingly W. Malesian in species diversity.

1. Monotoideae Gilg - Rays usu. uniseriate; adaxial gland at base of lamina; androgynophore +, exostome prolonged. - 3/30. Africa, Madagascar, South America.

2. Pakaraimaeoideae Maguire, Ashton & de Zeeuw - Included phloem +; C imbricate, shorter than K. - 1/1. The Guianan Highlands, South America.

3. Dipterocarpoideae Burnett - Resin ducts pervasive in wood; at least lateral bundles leaving central cylinder well before they enter the leaf; anthers ± basifixed, pollen tricolpate; fruit usu. 1-seeded, usu. a nut, endocarp hairy, K enlarging unequally and thickish; endosperm 0. - 13/650. Seychelles, Sri Lanka, India, South East Asia to New Guinea, but mostly W. Malesian.

Cytinaceae + Muntingiaceae: ellagitannins +; fruit a berry.

CYTINACEAE A. Richard - Echlorophyllous root parasites; vessels 0; inflorescence racemose; flowers imperfect; P 4-9, basally connate; staminate flowers: A 6-10, extrorse, connate, monothecate, nectariferous cavities between the stamens; carpellate flowers: nectary near base of style, G [8-14], inferior, placentation intrusive parietal, many atropous ovules/carpel, nucellar epidermis persists, micropyle endostomal; exotegmic cells thickened all around; embryo undifferentiated. - 2/10. Central America, Mediterranean, South Africa and Madagascar.

MUNTINGACEAE C. Bayer, M. W. Chase & M. F. Fay - Prophylls basal on axillary shoot, heteromorphic, stipule-like, stipules ?0; inflorescence extra-axillary, fasciculate; C crumpled in bud. - 3/3. Tropical America.

MALVACEAE Jussieu - Inflorescence made up of modified cymose units ["bicolor units"]; A fundamentally obdiplostemonous. - 243/4225+. Largely tropical, also temperate.

Grewioideae + Byttnerioideae: ?

1. Grewioideae Hochreutiner - 25/770. Pantropical (warm temperate).

2. Byttnerioideae Burnett - C clawed. - 26/650. Pantropical, esp. South America.

Sterculioideae + Tilioideae + Dombeyoideae + Brownlowioideae + Helicteroideae + [Malvoideae + Bombacoideae]: 21 bp deletion in ndhF gene.

3. Sterculioideae Burnett - Plant monoecious, inflorescence axillary, paniculate, lacking cymose units, epicalyx 0; C 0, androgynophore +, filaments connate, G largely free, micropyle endostomal; fruit a follicle. - 12/430. Pantropical.

4. Tilioideae Arnott - 2(?3)/50. N. temperate.

5. Dombeyoideae Beilschmied - 21/381. Old World tropics, St Helena, esp. Madagascar and Mascarenes.

6. Brownlowioideae Burret - 8/68. Tropical, esp. Old World.

7. Helicteroideae (Schott & Endlicher) Meisner - 8-10/95. Tropical, esp. Southeast Asia and W. Malesia.

Malvoideae + Bombacoideae: K connate, adaxially at base with multicellular clavate nectariferous hairs, C adnate to base of A, staminal tube +, anthers monothecal.

8. Malvoideae Burnett - 78/1670. Temperate to tropical.

9. Bombacoideae Burnett - Trunk often stout, with parenchymatous water-storage tissue, bark thin; leaves palmate; filaments usu. fasciculate, pollen ± flattened, triangular, nucellar cap +; cotyledons folded. - 16/120. Tropical, esp. America and Africa.


SANTALALES Dumortier  

Mycorrhizae absent; polyacetylenes [triglycerides with C18 acteylenic acids], triterpenic sapogenins +; perforation plates not bordered; intervascular pits alternate; pericyclic fibers 0; petiole/mesophyll with astrosclereids; lamina entire; inflorescences cymose; K often reduced, C valvate, A opposite C, anthers basifixed, pollen colpate or porate, disc +, 1 pendulous apotropous tenuinucellate ovule/carpel, micropyle bistomal, style single, stigma small, globose; fruit a drupe, 1-seeded, K persistent; seed coat crushed; endosperm cellular, chalazal haustorium +, embryo small/minute. - 8 families, 151 genera, 985 species.

ERYTHROPALACEAE - 13/65. Pantropical, SE Asia, but not Madagascar or New Guinea and to the SE.

"Olacaceae" [[[Misodendraceae + Schoepfiaceae] Loranthaceae] [Opiliaceae + Santalaceae]]: root parasites; vessel elements with simple perforations; rays >7 cells wide; nodes 1:1; sclerenchyma fibres of petiole and median vein 0; petiole bundle arcuate; stomata paracytic; silicification of some foliar mesophyll/epidermal cells; embryo sac with chalazal caecum and ± developed micropylar prolongation.

"OLACACEAE" R. Brown - 14/103. Pantropical (subtropics).

1. Olacoideae - 3/57. Pantropical (SE China, Formosa).

2. Anacalosoideae - 7/32. Pantropical (SE China, Formosa).

3. Ximenioideae - 4/13. Pantropical, warm temperate.

[[Misodendraceae + Schoepfiaceae] Loranthaceae] [Opiliaceae + Santalaceae]: G not septate, ovules ategmic; testa 0.

[Misodendraceae + Schoepfiaceae] Loranthaceae: cambium storied; petiole astrosclereids 0; epidermal cells sclerified, with druses; K minute, G [3].

Misodendraceae + Schoepfiaceae: ?

MISODENDRACEAE J. Agardh - Stem parasites; sieve tube plastids lacking starch and protein inclusions; stem apex aborting; plant dioecious; vegetative and reproductive shoots sharply distinguished; staminate flowers without a perianth; fruit a nut, with three persistent plumose staminodes. - 1/11. Cool temperate South America.

SCHOEPFIACEAE Blume - Bract immediately below and surrounding G; flowers distylous, C tubular, with adaxial hairs. - 3/55. Central and South America, a few species in tropical South East Asia-Malesia.

LORANTHACEAE Jussieu - Stomatal orientation transverse to long axis; collenchymatous zone below the embryo sacs; fruit with viscin; endosperm compound. - 68/950. World-wide.

1. Nuytsia - Fruit dry, winged; cotyledons 3-4, foliaceous. - 1/1. S.W. Australia.

2. The rest - Stem parasites, forming burl at point of attachment and with epicortical roots running over the surface and often forming secondary burls; embryo ± plug-shaped, no radicle. - 67/950. World-wide.

Opiliaceae + Santalaceae: P only.

OPILIACEAE Valeton - G superior. - 10/72. Pantropical, but Agonandra the only genus in America.

hen young; there is only one perianth whorl. The ovary is superior.

SANTALACEAE R. Brown - Cuticular epithelium +; stomatal orientation transverse to long axis. - 44/935. World-wide, esp. tropics.

BALANOPHORACEAE L. C. & A. Richard - Echlorophyllous root parasites from underground tuber-like structures, these rupture and leave a collar-like structure at ground level; roots 0; stems endogenous; inflorescence racemose, terminal; flowers small; A extrorse, usu. connate; carpellate flowers: P 0 or minute; embryo small to undifferentiated. - 7/50. Mostly tropical.



Tension wood?; crystals +; petiole bundle annular; stomata cyclocytic; filaments stout; style +; seed endotestal; endosperm development?, embryo? - 2 families, 3 genera, 4 species.

AEXTOXICACEAE Engler & Gilg - Pith heterogeneous; peltate scales +; bracteoles enveloping flower; K thin, deciduous, A 5, nectaries staminodial; fruit a drupe; seeds ruminate; embryo transverse. - 1/1. Chile.

BERBERIDOPSIDACEAE Takhtajan - Filaments short, anthers inserted along connective, connective with apical prolongation, placentation parietal; fruit a berry. - 2/3. Chile, E. Australia.

CARYOPHYLLALES + ASTERIDS: seed exotestal; embryo long.


Not mycorrhizal; root hair cells in vertical files; only alternate vascular pitting; perforation plates not bordered; leaves entire; anther with outer parietal cells developing directly into the endothecium, pollen colpate, spinulose, G [3], when G = K or P, opposite them, micropyle endostomal, styles/style branches long; fruit a loculicidal capsule. - 31 families, 692 genera, 11155 species.

Droseraceae [Tamaricaceae + Polygonaceae groups]: acetogenic napthoquinones [plumbagin and related compounds] +; endosperm starchy.

Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]]]: plumbagin +; plants carnivorous; vascularised multicellular glands; inflorescence cymose; C contorted, A extrorse, ovary unilocular.

DROSERACEAE Salisbury - Herbs; leaves adaxially circinate, with glandular hairs, sensitive; pollen in tetrads, porate, placentation parietal. - 3/115. World-wide.

Nepenthaceae [Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]]: rays 1-2 cells wide; cortical bundles in stem; petiole bundle(s) surrounded by massive sclerenchymatous ring with embedded vascular bundles; anthers basifixed.

NEPENTHACEAE Berchtold & J. Presl - Climbers; large spirally-thickened cells in pith, cortex, etc.; leaves with foliaceous basal part, a narrow twining portion, and pitcher-like terminal portion covered by a lid; plant dioecious; inflorescence racemose; P uniseriate, with nectariferous glands adaxially, A connate into a central column, pollen in tetrads, apertures indistinct, placentation axile, style short, stigma broad. - 1/90. Madagascar to New Caledonia.

Drosophyllaceae [Ancistrocladaceae + Dioncophyllaceae]: ?

DROSOPHYLLACEAE Chrtek, Slavíkovà & Studnicka - Leaves abaxially circinate, with glandular hairs; pollen polyporate; placentation basal. - 1/1. Iberian Peninsula, Morocco.

Ancistrocladaceae + Dioncophyllaceae: lianes; (acetogenic isoquinoline alkaloids +); cortical bundles 0; petiole with inverted bundles in sclerenchyma ring; stomata actinocyclic; anthers introrse.

ANCISTROCLADACEAE Walpers - Not carnivorous; modified branch systems with grapnels in series; lamina with glands in pits on abaxial surface; G half or more inferior, with 1 basal hemitropous ovule; fruit a nut, K much enlarged; seed ruminate. - 1/12. Tropical Africa to W. Borneo and Formosa.

DIONCOPHYLLACEAE Airy Shaw - Plant with rosette leaves first, venation closely parallel, then leaves with a short blade and glandular hairs on the abaxial surface of the prolonged midrib, then leaves with paired grapnel hooks at the apex; placentation parietal; fruits open early; seeds flattened, on elongated funicles. - 3/3. Tropical West Africa.

Frankeniaceae + Tamaricaceae + Plumbaginaceae + Polygonaceae: sulphated flavonols, ellagic acid +; seed exotestal.

Frankeniaceae + Tamaricaceae: halophytic; bisulphated flavonols +, myricetin 0; wood storied; ?silica bodies; leaves small, with salt-excreting glands; flowers small, 4-6-merous, C with basal adaxial appendages, pollen not spinulose, placentation (basal; intrusive-)parietal; exotestal cells bulging or as hairs; endosperm +.

FRANKENIACEAE Desvaux - Leaves opposite. - 1/90. ± World-wide in warm, dry areas, but scattered.

TAMARICACEAE Berchtold & J. Presl - Embryo sac tetrasporic. - 5/90. Eurasia and Africa, esp. Mediterranean to Central Asia.

Plumbaginaceae + Polygonaceae: O-methylflavonols, myricetin, quinones +; (wood storeyed); successive cambia 0; cortical and/or medullary vascular bundles +; nodes 3:3; leaf bases broad; pollen grains usu. with starch, G with median member adaxial, 1 loculus and 1 basal ovule; fruit surrounded by accrescent calyx which forms part of the dispersal unit; exotesta ± persistent, otherwise seed coat undistinguished.

PLUMBAGINACEAE Jussieu - A opposite C, G [5], ovule circinotropous, funicle long and curved, embryo sac tetrasporic, 4- or 8-nucleate. - 27/836. Predominantly Mediterranean to Central Asia, scattered elsewhere.

1. Plumbaginoideae Burnett - Capsule basally circumscissile. - 4/36. East Asia and Africa.

2. Staticoideae Kostel. - C connate, A adnate to C.

2A. Aegilitideae (Link) Peng - Ellagic acid +; successive cambia +; cortical vascular bundles +. - 1/2. Malesian mangroves.

2B. Staticeae Bartling - Leaves more or less clustered at base; inflorescence leaves reduced or absent; inflorescence axis channelled; K scarious; deletion of rpl16 intron. - 22/800. Mostly Irano-Turanian (Mediterranean), but also S. Africa, S. South America, and W. Australia.

POLYGONACEAE Jussieu - Pits vestured; stipule sheathing the stem; P 5, ovule atropous; achene angled. - 43/1100. World-wide.

1. Polygonoideae Jaretzky - 15/590. Especially (warm) temperate.

2. Eriogonoideae Arnott - 28/520. Largely tropical, especially America, Eriogonum and relatives esp. W. North America.

Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]]]: stylodia stigmatic their entire length; endosperm slight.

RHABDODENDRACEAE Prance - Sieve tube plastids with protein crystalloids and starch; nodes multilacunar; secretory cavities with resin; foliar fibre-like sclereids +; hairs peltate, fringed; leaves revolute, obscurely punctate; A many, filaments very short, anthers very long, pollen not spinose, G 1, with 1 (2) basal campylotropous ovules, bitegmic zone short, archesporium multicellular, style basal; fruit a drupe, pedicel apex swollen. - 1/3. Tropical South America.

Simmondsiaceae [[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]]: nodes 1:1; C 0.

SIMMONDSIACEAE van Tieghem - Petiole articulated; staminate plant: A extrorse, anthers longer than connective, pollen ± porate, central part operculoid; carpellate plant: flowers axillary, 1 apical pendulous apotropous ovule/carpel, outer integument ca 10 cells and inner integument 4-5 cells thick, styles papillate all around; fruit with persistent columella; seed 1, testa multiplicative, vascularised, exotestal cells palisade, walls thickened. - 1/1. S.W. North America.

[Asteropeiaceae + Physenaceae] [Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.]: ?

Asteropeiaceae + Physenaceae: successive cambia 0; fruit single-seeded.

ASTEROPEIACEAE Reveal & Hoogland - Pedicels with many bracteoles; C +, deciduous, stigma continuous across G; cotyledons spirally twisted. - 1/8. Madagascar.

PHYSENACEAE Takhtajan - Plant dioecious; anthers long; seed large, coat vascularised, 16-20 layers thick, walls not notably thickened. - 1/2. Madagascar.

Caryophyllaceae, Nyctaginaceae, Cactaceae, etc., "Core Caryophyllales": Plant herbaceous; CAM [especially pervasive in succulents] and C4 photosynthesis common; ferulic acid bound to unlignified cell walls; (O-methylated) flavonols, quinones, betalains [chromoalkaloids], triterpenoid saponins +, tannins, myricetin 0; (phytoferritin +); sieve tube plastids with a ring of proteinaceous filaments and a central angular crystalloid; pericyclic fibers 0; inflorescence cymose; pollen trinucleate, (polyaperturate), foot layer thin, nectaries on adaxial bases of stamens, ovary 1-locular, ovules campylotropous, (funicles long), stigmas papillate, little expanded; seeds with exotestal and endotegmic cells thickened, bar-like thickenings on endotegmic cells; endosperm 0, perisperm +, starch grains clustered, embryo curved; mitochondrial rps 10 gene and chloroplast rpl2 intron lost.

Caryophyllaceae + Achatocarpaceae + Amaranthaceae: mitochondrial rps1 and 19 genes lost .

CARYOPHYLLACEAE Jussieu - Anthocyanins +; "C" present; endotegmen lacking bars. - 86/2200. Mostly temperate.

Achatocarpaceae + Amaranthaceae: ?

ACHATOCARPACEAE Heimerl - 3/7. S.W. U.S.A. to South America.

AMARANTHACEAE Jussieu - Sieve tube plastids lacking protein crystalloid; pollen wall foot layer well developed. - 174/2050. ± World-wide, esp. warm and dry temperate and subtropics, esp. saline habitats.

Stegnospermataceae [Limeaceae [[Lophiocarpaceae [Aizoaceae, Nyctaginaceae, etc.]] [Molluginaceae, Cactaceae, etc.]]: ?

STEGNOSPERMATACEAE Nakai - "C" (2-)5, 1 basal ovule/carpel; seeds arillate. - 1/3. Central America, the Antilles.

Limeaceae [[Lophiocarpaceae [Aizoaceae [Giseckiaceae [Nyctaginaceae, etc.]]]] [Molluginaceae, Cactaceae, etc.]]: ovules apotropous.

LIMEACEAE Shipunov - Anthocyanins +. - 2/23: Southern Africa, to Ethiopia, S. Asia, also Australia.

[Corbichonia + Lophiocarpus] + Aizoaceae, Nyctaginaceae, etc. + Molluginaceae, Cactaceae, etc.: sieve tube plastids with globular crystalloids

[Lophiocarpaceae [Aizoaceae [Giseckiaceae [Nyctaginaceae, etc.]]]]: ?

LOPHIOCARPACEAE Doweld & Reveal - 2/6. Africa, esp. SW Africa.

Barbeuiaceae [Aizoaceae [Gisekiaceae [Sarcobataceae + Phytolaccaceae + Nyctaginaceae]]]: ?

BARBEUIACEAE Nakai - Sieve tube plastids with polygonal crystalloids. - 1/1. Madagascar.

Aizoaceae [Sarcobataceae + Phytolaccaceae + Nyctaginaceae]: soluble oxalate accumulation; raphides +.

AIZOACEAE Martynov - Wood rayless; leaf trace bundles forming reticulum in cortex; epidermis with bladder-like cells; hypanthium +, P colored internally, often with subapical abaxial appendage ["horn"], A centrifugal, many, primordia 5, nectary as ring, G septate. - 123/ca 2020. Mostly southern Africa, also Australia, etc., tropical and subtropical, arid.

1. Sesuvioideae Lindley - Stipules +; prophylls often prominent; inflorescence usu. distinct from vegetative plant; A primordia opposite P; capsule circumscissile; seeds arillate. - 5/36. Tropics and Subtropics.

Aizooideae [Mesembryanthemoideae + Ruschioideae]: A primordia alternate with P; fruit a hygrochastic capsule.

2. Aizooideae - Bladder hairs with large terminal cell and multicellular stalk[?]; inflorescence leafy; seeds upright [?]. 7/135. Drier parts of S. Africa, also Australia (Gunniopsis), few N. Africa and Asia Minor, N. America, etc.

Mesembryanthemoideae + Ruschioideae: leaves very succulent; "C" [staminodial] many, G more or less inferior, nectary interrupted; x = 9.

3. Mesembryanthemoideae Ihlenfeldt, Schwantes & Straka - Cortical bundles +; inflorescence not distinct; K, "C" and A basally connate, nectary hollow or shell-shaped ["koilomorphic"], placentation axile. - 12/80. S. Africa, Mesembryanthemum also W. North and South America, Australia, N. Africa, the Mediterranean and the Near East.

4. Ruschioideae Schwantes - Bladder-like epidermal cells 0; inflorescence distinct; filaments papillate, placentation basal or parietal; capsule with expanding keels more or less restricted to valves. - Ca 110/ca 1765. Southern Africa, esp. the western coastal Succulent Karroo.

Giseckiaceae [Sarcobataceae + Phytolaccaceae + Nyctaginaceae]: 1 basal ovule/carpel; ORF 2280 sequence similarity, 210 bp deletion in chloroplast genome.

GISECKIACEAE Nakai - Anthocyanins +; raphides +; inflorescence leaf-opposed; G pseudapocarpous; fruit muricate. - 1/5. Africa, Asia.

Sarcobataceae + Phytolaccaceae + Nyctaginaceae: ?

Sarcobataceae + Phytolaccaceae + Nyctaginaceae: 210 bp deletion in chloroplast.

SARCOBATACEAE Behnke - Bracteoles 0; staminate inflorescence catkinate, flowers with peltate scales ["bracts"], anthers long, pollen pantoporate, pore margins raised; carpellate flowers single, P tubular, bilobed [?bracteoles], G [2], ovule 1. - 1/2. S.W. North America.

Phytolaccaceae + Nyctaginaceae: 1 basal ovule/carpel; protein bodies in nuclei.

PHYTOLACCACEAE R. Brown - Inflorescence racemose; fruit a berry. - 18/65. Tropical and warm temperate, esp. America.

1. Phytolaccoideae - 4/31. Chile, Mexico, or cosmopolitan (Phytolacca).

2. Rivinioideae Nowicke - 9/13. Central and South America, Antilles, Florida, tropical Africa (some Hilleria), Australia, New Hebrides and New Caledonia (Monococcus).

3. Agdestidoideae Nowicke - 1/1. S. U.S.A. to Niacaragua.

NYCTAGINACEAE A.-L. de Jussieu - P connate, petaloid, lobes induplicate-valvate or contorted, G 1, ovule 1, basal, funicle short; fruit achene or nutlet. - 30/395. Tropical to warm temperate.

1. Leucastereae Bentham & Hooker - Indumentum ± stellate; style thick/0, stigma crest-like. - 4/5. S.E. South America, esp. Brasil.

Boldoeae + The Rest: style long, slender.

2. Boldoeae Heimerl - Stigma inconspicuous. - >3/3. Mexico to Bolivia, the West Indies.

3. The rest - Leaves opposite; P bipartite, tube stout, limb thin, pollen pantoporate; fruit with accrescent basal part of P tube. - 25/390. Tropical to warm temperate, esp. herbs and shrubs in arid southwestern North America and arborescent genera in the Neotropics and Caribbean.

Molluginaceae [Halophytaceae + Didieraceae + Basellaceae + Montiaceae [Talinaceae [Portulacaceae [Anacampseros, etc. + Cactaceae]]]]: ?

MOLLUGINACEAE Rafinesque - Anthocyanins, hopane saponins +; leaves pseudoverticillate. - 9/87. Largely S. Africa, a few ± tropical to warm temperate.

Halophytaceae + Didieraceae + Basellaceae + Montiaceae [Talinaceae [Portulacaceae [Anacampseros, etc. + Cactaceae]]]>: plants succulent; CAM +; secondary growth normal; phloem parenchyma cells with phytoferritin; Ca oxalate crystals in stem epidermis; P petaloid, pollen with similar fused columellae, etc.

HALOPHYTACEAE A. Soriano - Wood rays 0; staminate flowers: inflorescence densely spicate, P 4, valvate-decussate, A extrorse, porose, anther dehiscence by contraction of the connective, endothecium with frame-shaped thickening on anticlinal walls, pollen cuboid, hexaporate; carpellate flowers: inflorescence fasciculate, P 0, G unilocular, with 1 basal ovule; fruit a nutlet, several becoming enclosed by inflorescence axis. - 1/1. Argentina.

Didieraceae + Basellaceae + Montiaceae [Talinaceae [Portulacaceae [Anacampseros, etc. + Cactaceae]]]: median bracteoles [lacking subtending buds] present, enclosing flower; median P abaxial [opposite outer median bracteole], pollen pantocolpate, G 3, no funicular obturator; 6-bp deletion in ndhf gene.

Didiereaceae + Basellaceae: stomata paracytic; ovary with single basal ovule; fruit indehiscent.

DIDIEREACEAE Radlkofer - 7/16. Madagascar, South Africa, E. Africa.

BASELLACEAE Rafinesque - Vines with swollen rhizomes or tubers; successive cambia +; inflorescence racemose; pollen hexacolpate/porate. - 4/19. Pantropical, but mostly New World.

Montiaceae [Talinaceae [Portulacaceae [Anacampseros, etc. + Cactaceae]]]: stomata parallelocytic.

MONTIACEAE Rafinesque - Annual to perennial herbs, often with swollen roots and basal leaves, or subshrubs; leaves often with clasping bases; outer wall of exotesta thickened and with stalactite-like projections. - ca 10/: Especially Western North and South America, also the Antilles and the Subantarctic Islands.

Talinaceae [Portulacaceae [Anacampseros, etc. + Cactaceae]]: ± tuberous; pericarp 2-layered, exocarp ± caducous.

TALINACEAE Doweld - Shrubs; leaves with paired axillary scales; pericarp epidermis papillate. - 2/22. America and Africa.

Portulacaceae [Anacampseros, etc. + Cactaceae]: cork cambium cortical; loss of pericyclic fibers; sclereids in stem cortex; leaves with axillary uni- or biseriate hairs/scales +.

PORTULACACEAE Jussieu - Succulent herbs; leaf stomata transverse; leaves ± terete; capsule circumscissile, pericarp undifferentiated. - 1/40-100. Worldwide, but especially tropical and subtropical North and South America.

Anacampseros, etc. + Cactaceae: A many.

Anacampseros etc. - Subshrubs; leaf stomata transverse; leaves ± terete; stigma receptive on both surfaces; exotesta ± separate, thin walled, unlignified; embryo slightly curved, not surrounding poorly developed perisperm. - 3-8/32. C. and S. Australia, Somalia to South Africa (most species), S. South America, N. Mexico and S.W. U.S.A.

CACTACEAE Jussieu - Short shoots producing spines and hairs; median bracteoles 0; P several-numerous, spiral, sepaline outside and petaline inside, G inferior, placentation parietal; fruit, and also funicles, fleshy. - 100/1500. Mostly New World, esp. arid conditions, but also rain forest climbers and epiphytes.

1. Rhodocactus - 1/7. Mexico and the Caribean, Brasil.

Pereskioideae [Maihuenioideae [Opuntioideae + Cactoideae]]: delayed bark formation; stomata in stem epidermis.

2. Pereskioideae Engelmann - 1/9. Andean, S. South America.

Maihuenioideae [Opuntioideae + Cactoideae]: stems succulent, wide band tracheids in secondary xylem with annular thickenings; hypodermis collenchymatous, intercellular spaces +, cortical chlorenchyma forming mesophyllar tissue; wide-band tracheids +; inflorescences axillary, flowers single; hypanthium +.

Maihuenioideae + Opuntioideae: Leaves terete.

3. Maihuenioideae Fearn - Bark formation not delayed; stem stomata 0; leaves with cylindrical reticulum of bundles, the external xylem surrounding central mucilage reservoir. - 1/2. Argentina and Chile.

4. Opuntioideae Burnett - Glochids with retrorse barbs; subsidiary cells of stomata do not, or only barely, overlap the ends of the guard cells; leaves ± terete; pollen porate; seeds surrounded by bony funicular "aril". - 5/225. S. Canada to Patagonia.

5. Cactoideae Eaton - Plant leafless [leaves up to 1.5(-2.5) mm long when mature]; calcium oxalate often as weddellite [CaC2O4.2H2O]; pollen colpate; testa interstitially pitted or cratered, outer periclinal wall much thickened; rpoC1 intron lost. - 92/1250. New World, (S. Canada to S. W. U.S.A. southwards. Rhipsalis also with a few (?native) spp. in Africa and Madagascar.

ASTERIDS - Sympetalae redux?: Nicotinic acid metabolised to its arabinosides; (iridoids +); C sympetalous, anthers dorsifixed?, (nectary gynoecial), ovules unitegmic, integument thick, endothelium +, nucellar epidermis does not persist, style single, long; seed exotestal; endosperm cellular, embryo long.

CORNALES Dumortier

Iridoids diverse, ellagic acid +, flavones 0; vessel elements with scalariform perforations; nodes 3:3; inflorescence cymose; C valvate, apparently free, tube formation early, A basifixed, G inferior, disc +, ventral carpellary bundles in the carpel wall [transseptal bundles, i.e. vascular bundles to ovules go over the top of the septum and then down; there are no bundles running up the central axis of the gynoecium], 1-2 apical crassinucellate ovules/carpel; fruit drupaceous, cells of wall sclereidal, germination valve(s) in the stone, K persistent. - 7 families, 51 genera, 590 species.

Cornaceae + Nyssaceae: route I secoiridoids, triterpenoid saponins +; hairs T-shaped, unicellular; K small, pollen with complex endaperture [a pore joining two lateral thinnings parallel to the colpus], style short.

CORNACEAE Dumortier - 2/85. Scattered, not S. South America.

NYSSACEAE Dumortier - Cells of fruit wall fibrous. - 5/22. Mainly East Asia, also Indo-Malesia and E. North America.

Hydrangeaceae + Loasaceae: similar route I secoiridoids and route II decarboxylated iridoids [e.g. deutzioside], flavonols +, ellagic acid 0; cork deep-seated; hairs tuberculate, walls calcified, with basal cell pedestals; leaves opposite, with glandular teeth (lobed); A 2x C, gynoecium with axial/central vascular bundles, placentation parietal, many ovules/carpel, stigma dry; fruit septicidal; exotestal cells variously elongated, inner walls thickened; mitochondrial coxII.i3 intron 0.

HYDRANGEACEAE Dumortier - 17/190. Warm temperate, a few tropical.

1. Jamesioideae Hufford - Leaf buttresses conspicuous after leaf fall; K valvate, C free, style branched. - 2/ca 5. W. North America.

2. Hydrangeoideae Burnett - Raphide sacs +. - 15/185. Warm temperate, esp. South East Asia and North America, S. to Chile and Malesia.

2a. Philadelpheae - Embryo sac ± protruding from the nucellus. - 6/130. Warm temperate, esp. South East Asia to the Philppines, S.W. North America, also Central America, one sp. in Europe.

2b. Hydrangeeae - C valvate, not clawed; fruits loculicidal, (baccate). - 9/65. Warm N. temperate, S. to Chile and Malesia.

LOASACEAE Jussieu - Vessel elements with simple perforations; trichomes glochidiate, often silicified; integument very thick, style hollow. - 14/265. Mostly American, also Africa and the Marquesas Islands.

1. Eucnide - A initiation centripetal, connate basally; n = (?19-)21. - 1/15. S.W. North America.

2. Schismocarpus - Filaments shorter than the anthers. - 1/1. Mexico.

Loasoideae + Mentzelioideae + Gronovioideae: G 3-5, when 3, odd member adaxial.

3. Loasoideae - Petals cymbiform, clawed, stamens in groups opposite C, pollen ?not striate, antesepalous staminodes + [outer whorl connate, as scales, inner whorl separate, more elaborated]; n = 6. - America, but also Africa and the Marquesas Islands.

Mentzelioideae + Gronovioideae: loss of C-A synorganisation.

4. Mentzelioideae - A centripetal, connate basally; n = 7. - 1/60. America.

5. Gronovoioideae - C valvate, petals with a single vascular trace, A 5, opposite K, anthers bifacial, G with 1 apical crassinucellate ovule, funicular obturator +; fruit a cypsela; testa none; endosperm haustoria 0. - America.

HYDROSTACHYACEAE Engler - Submerged herbs; primary root 0; stomata 0; leaves in a rosette, deeply and complexly divided, surface with small enations, stipule +; inflorescence spicate, plants di(mon)oecious, P 0, nectary 0; staminate flowers: A 2, monothecal, extrose, pollen in tetrads, inaperturate; carpellate flowers: G [2], transverse, placentation parietal, many unitegmic tenuinucellate ovules/carpel, styles separate, filiform; fruit a septicidal capsule; seeds minute, outer cell walls much thickened, mucilaginous; endosperm scanty or 0, micropylar haustorium +. - 1/20. C. and S. Africa, Madagascar.

Grubbiaceae + Curtisiaceae: leaves opposite, bases connected by a ridge; flowers small, one ovule/carpel, style short; endosperm copious.

GRUBBIACEAE Endlicher - Ericoid shrublet; anthers bisporangiate/unithecal, ?inverted; fruit a syncarp. - 1/3. Cape Province, South Africa.

CURTISIACEAE Takhtajan - Gynoecium with axial/central vascular bundles. - 1/2. S. Africa.

ERICALES + EUASTERIDS: ovules tenuinucellate.

ERICALES Dumortier

Nodes 1:1; leaves spiral, teeth with single vein and opaque deciduous cap. - 25 families, 346 genera, 11,515 species.

Balsaminaceae + Marcgraviaceae + Tetrameristaceae: myricetin +, ellagic acid 0; vessel elements with simple perforations; raphide sacs +, druses 0; lamina toothed; inflorescence racemose; A = and opposite K, anthers (near) basifixed, nectary 0, ovules bitegmic, micropyle endostomal, style short, stigma little expanded; endosperm with micropylar haustorium.

BALSAMINACEAE Berchtold & J. Presl - Herbaceous; flowers vertically monosymmetric, inverting during growth; functionally abaxial sepal with prominent spur, nectary inside, adaxial C with a sepaloid keel, lateral petals connate in pairs, A forming cap over stigma, filaments stout, partly connate apically, anthers connate and forming cap over stigma, pollen with cellulose threads holding it to anther, grains colpate, integuments intermediate unitegmic-bitegmic [with a degree of congenital fusion]; fruit dehiscence along the septal radius; seed pachychalazal. - 2/1000. Mostly Old World, Africa (esp. Madagascar) to mountains of SE Asia.

Marcgraviaceae + Tetrameristaceae: vessels in radial multiples; ± branched sclereids +; lamina supervolute, elongating in bud, with obscure abaxial lines; fruit indehiscent.

MARCGRAVIACEAE Candolle - Stomata staurocytic; lamina entire, with marginal to abaxial cavities; inflorescence a raceme, bracts abaxially ascidiate, nectariferous; C ± connate; fruit with fleshy placentae; seeds many, small. - 7/130. New World tropics.

TETRAMERISTACEAE Hutchinson - Petioles short; bracteoles rather large, ± caducous; K with adaxial glands, 1 ovule/carpel. - 3/5. W. Malesia, Central and N. South America.

Other Ericales: C tube well developed, style long.

Polemoniaceae + Fouquieriaceae: cork outer cortical; vessel elements with simple perforations; C connate, nectary +, G [3], style strongly lobed.

POLEMONIACEAE Jussieu - K connate, A = and opposite K, adnate to C. - 18/385: N. temperate, W. North America, South America.

1. Polemonioideae Arnott - 13-22/350. W. North America, N. temperate, southern South America.

2. Cobaeoideae Arnott - K not differentiated; seeds winged. - 5/35. Baja California, tropical America.

3. Acanthogilioideae J. M. Porter & L. A. Johnson - Leaves very dimorphic, persistent branched spines on long shoots, deciduous and unlobed on short shoots; seed winged. - 1/1. Baja California.

FOUQUIERACEAE Candolle - Xeromorphic, with long and short shoots; leaves heteromorphic, spines developed from petioles; seeds winged. - 1/11. S.W. North America.

LECYTHIDACEAE A. Richard - Cortical bundles +; A many, G inferior, ovules bitegmic, micropyle endostomal, archesporium multicellular, style 0/short; testa multiplicative, ± lignified; endosperm nuclear, 0. - Ca 25/310. Tropical, especially America and W. Africa.

1. Napoleonaeoideae Niedenzu - A extrorse, style 0, stigma broad, pentagonal, flat. - 2/11. W. tropical Africa.

Scytopetaloideae + Planchonioideae + Foetidioideae + Lecythidoideae: ?

2. Scytopetaloideae Appel - Pollen tricolpate or tricolporoidate, G superior (half superior), endothelium +, style relatively long, slender; endosperm ruminate, 0. - 6/21. Africa, South America.

Planchonioideae + Foetidioideae + Lecythidoideae: ?

Planchonioideae + Foetidioideae: cortical bundles inverted; fruit indehiscent.

3. Planchonioideae Niedenzu - A with a basal ring, pollen syncolpate, strong colpus margin ridge. - 6/58. Paleotropical.

4. Foetidioideae Niedenzu - SiO2 bodies +; C 0, placentae peltate. - 1/17. E. Africa to Mauritius.

5. Lecythidoideae Niedenzu - Staminal ring +. - 10/200. Neotropical.

Sladeniaceae + Pentaphylacaceae: mucilage cells +; hairs unicellular; C ± campanulate, only basally connate, fairly small [petals 1> cm long], A basifixed, pollen 14-30 µm long, surface usu. little ornamented, nectary 0, placentae becoming ± swollen; fruit a capsule, columella persistent; endosperm +.

SLADENIACEAE Airy Shaw - Inflorescence cymose, branches widely spreading; flowers small, A opening apically. - 2/3. S.E. Asia, tropical E. Africa.

PENTAPHYLACACEAE Engler - Inflorescence of axillary flowers or fasciculate; mesotesta well developed; embryo U-shaped. - 12/337. Tropical and subtropical, but few in Africa.

1. Pentaphylaceae - A 5, filaments very broad, narrowed and incurved apically, theacae each opening by a valve that lifts up, 2 apical ovules/carpel; midrib ["teeth"] separating from rest of valves; cotyledons longer than radicle. - 1/1. Kwangtung and Hainan to Sumatra, scattered.

Ternstroemieae + Frezierieae: Ellagic acid +; sclereids +; perulae 0; filaments to 2x longer than anthers, connective usu. prolonged; fruit ± fleshy; mesotesta lignified, ± crystalliferous; endosperm +, embryo incumbent, radicle longer than cotyledons.

2. Ternstroemieae Candolle - Sclereids much branched; leaves pseudoverticillate; flowers single from axils of reduced leaves; K opposite C, filaments shorter than anthers, 4-12 apical ovules/carpel; fruit irregularly dehiscing; seeds few, 3< mm long, brown, sarcotestal, exotesta 10< cells across, sclerified mesotesta 7-15 cells across. - 2/103. Tropics, esp. Malesia and Central to South America, in W. Africa only 2 species.

3. Frezierieae - Sclereids little branched; 4-many ovules/carpel; fruit a berry; seeds many, <4(-6) mm long, brown or black, sclerified mesotesta 1-5 cells across. - 9/233. SE Asia to Malesia, Hawaii, Central to South America, E. Africa (Balthasaria) and Canaries (Visnea).

Sapotaceae [Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]: ellagic acid 0; C connate.

SAPOTACEAE Jussieu - Plant with gutta; wood siliceous and/or with SiO2 grains; T-shaped hairs +; A = and opposite C, G hairy; fruit a berry; seed coat hard, shiny, with large hilum. - 53/975. Pantropical.

1. Sarcospermatoideae (Lam) Swenson & Anderberg - Leaves ± opposite, stipules cauline; inflorescence axis apparently well developed [actually a reduced branch]; G 1-[2], style stout. - 1/6. IndoMalesian.

Sapotoideae + Chrysophylloideae: staminodes +, G [(2-)3-many].

2. Sapotoideae Eaton - Pantropical

3. Chrysophylloideae Luersson - Stipules 0; endosperm copious, cotyledons foliaceous. - 25/. Pantropical.

Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]: ?

EBENACEAE Gürcke - Leaves 2-ranked, margins entire, flat glands on lower surface; inflorescence cymose; C contorted, A 2x K, adnate to C, 2 pendulous ovules/carpel; fruit a berry; testa vascularized. - 5/548. Tropical (to temperate).

Lissocarpoideae Wallnöfer - Plant glabrous; pollen triporate; G inferior. - 1/8. Tropical South America.

2. Diospyroideae - 3/540. Tropical (to temperate).

Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]: schizogenous secretory canals +; nodes ?3:3; (stomata anisocytic); small ± immersed often peltate glandular hairs +; inflorescence racemose; C and A from common primordia, A = and opposite C, antesepalous whorl at least a vascular trace, nectary +, G [5], placentation free-central, ovules at least partly immersed in swollen placenta, bitegmic, micropyle bistomal, endothelium +, tanniniferous, style short, stigma ± capitate; seeds angled; endotesta crystalliferous; endosperm nuclear, copious, cell walls thick, amyloid or hemicellulosic.

MAESACEAE Anderberg, B. Ståhl & Kallersjö - Stamen primordium smaller than petal primordium, G inferior. - 1/150. Old World tropics to Japan, the Pacific, and Australia.

Theophrastaceae [Myrsinaceae + Primulaceae]: herbs[?]; rays >5-seriate, uniseriate rays 0; bracteoles 0; C imbricate, stamen primordium larger than petal primordium, subrotate, tube rather short.

THEOPHRASTACEAE Link - Bracts displaced up the pedicels; staminodes +, petaloid. - 5/105. Mostly New World and tropical, some also more temperate and Old World.

1. Samolus - G semi-inferior. - 1/15. America, the Antipodes, Europe, tropical to temperate.

2. The rest (Theophrastaceae s. str.) - Shrubs; anthers extrorse, with calcium oxalate. - 4/90. New World tropics.

Primulaceae + Myrsinaceae: two ndhF deletions.

PRIMULACEAE Borkhausen - Secretory canals 0; inflorescence scapose. - 9/900. Northern hemisphere, scattered elsewhere.

MYRSINACEAE R. Brown - 41/1435. Pantropical and N. Temperate.

MITRASTEMONACEAE Makino - Root parasites; leaves opposite, scale-like; flowers quite large, single; P 4, connate, A extrorse, completely connate and surrounding G except for small apical pore, polythecate, pollen 2-aperturate, G with 9-20 intrusive parietal placentae, ovules very numerous, with a funicular obturator, style stout, stigma hemispherical; fruit berry-like, circumscissile; funicle sticky; embryo unorganised, 4-celled. - 1/1. South East Asia to Malesia.

Theaceae + Symplocaceae, etc., + Ericaceae, etc.: ?

THEACEAE Ker Gawler - Ca 7/195(-460!). Mostly South East Asia-Malesia, also S.E. U.S.A.

1. Theeae Szyszylowicz - Pedicels multibracteolate, K and C intergrading, A in 2 whorls; capsule with columella; seeds winged or not. - 5/. Southeast Asia, Malesia, tropical America.

2. Gordonieae de Candolle - A in 3-5 whorls [?is this a character]; capsule with columella, dehiscence also septicidal; seeds apically winged (not – Franklinia]. - 3/. Southeast Asia, West Malesia, S.E. United States.

3. Stewartieae Choisy - Capsule lacking columella; seeds narrowly winged or not. - 1/9. East Asia, E. North America.

Symplocaceae [Styracaceae + Diapensiaceae]: inflorescence racemose; endosperm copious.

SYMPLOCACEAE Desfontaines - G inferior. - 1/320. Tropical to subtropical, inc. New Caledonia, not Africa.

Styracaceae + Diapensiaceae: cork pericyclic; glandular hairs 0; leaves spiral, margins toothed or entire; A basifixed, style continuous, hollow; fruit a capsule.

STYRACACEAE Candolle & Sprengel - Hairs stellate; K open, linear anther connectives of equal length throughout, ovules apotropous; hairs inside the carpel loculi. - 11/160. Warm N. temperate to tropical.

DIAPENSIACEAE Lindley - Herbs; secondary wood rays 0; anthers ± incurved, thecae ± horizontal. - 6/18. Arctic and N. temperate, esp. East Asia and E. U.S.A.

[Sarraceniaceae [Actinidiaceae + Roridulaceae]] [Clethraceae [Cyrillaceae + Ericaceae]]: anthers extrorse, inverting during development, opening by pores or short slits, tectum and foot layer solid, infratectum with granular elements, endothelium +, style impressed; exotesta with strongly thickened inner walls; endosperm copious.

Sarraceniaceae [Actinidiaceae + Roridulaceae]: route I secoiridoids +; nectary 0; hypostase +.

SARRACENIACEAE Dumortier - Insectivorous rosette herbs; leaves with pitchers; pollen 5+ colporoidate. - 3/ca 15. E. and W. U.S.A. and the Guyana Highlands.

Actinidiaceae + Roridulaceae: ?

ACTINIDIACEAE Gilg & Werdermann - Raphides +; seeds embedded in placental tissue. - 3/355. Largely tropical, esp. South East Asia to Malesia, but not Africa.

RORIDULACEAE Berchtold & J. Presl - Hairs capitate, secreting resin; leaves linear; A 5, endothecium 0. - 1/2. Southern Africa.

Clethraceae [Cyrillaceae + Ericaceae]: ellagic acid +; cork pericyclic; pericyclic fibers absent; bracteoles 0; flowers pendulous, A = 2x K, nectariferous disc +, style hollow; endosperm with micropylar and chalazal haustoria; embryo terete.

CLETHRACEAE Klotzsch - 2/76. E. Asia to Malesia, S.E. U.S.A., Mexico southwards on mountains, Cuba, 1 sp. on Madeira; largely tropical montane or subtropical.

Cyrillaceae + Ericaceae: myricetin +; colleters +; C connate, nectary at base of G, stigma wet.

CYRILLACEAE Endlicher - Sieve tube plastids with protein crystalloids and fibers; anthers introrse, not inverting, opneing by slits, style short; testa 0. - 2/2. S. U.S.A. to N. South America.

ERICACEAE Jussieu - Hair roots present [consisting of endodermis, exodermis, tracheid, sieve tube + companion cell], invested by hyphae [Hartig net]; leaf teeth associated with usually glandular headed multicellular hairs; A obdiplostemonous, inverting late. - Ca 126/3995. World-wide, but rare in lowland tropics.

Enkianthoideae Kron, Judd & Anderberg - Pith with small, thick-walled and lignified and larger and thin-walled cells mixed [heterogeneous]; leaves pseudoverticillate; anthers with paired awns; megagametophyte with "ears". - 1/16. South East Asia: China, Japan and environs.

Monotropoideae + Arbutoideae [[Cassiopoideae + Ericoideae] [Harrimanelloideae [Styphelioideae + Vaccinioideae]]]: fungal hyphae with complex coiled intrusions into the exodermal cells of hair roots [pegs only in Monotropeae, Pterosporeae]; anthers with exothecium; no chalazal vascular bundle in seed.

Monotropoideae Arnott - Herbaceous; C free. - 15/50. N. hemisphere, largely temperate.

Arbutoideae [[Cassiopoideae + Ericoideae] [Harrimanelloideae [Styphelioideae + Vaccinioideae]]]: (petiole bundle annular); bracteoles + [but see Monotropoideae], anthers without endothecium, porose, pollen in tetrads.

Arbutoideae Niedenzu - 1-6/ca 80. Warm (cold) temperate, esp. S.W. North America, Mediterranean.

Cassiopoideae + Ericoideae + Harrimanelloideae + Styphelioideae + Vaccinioideae: ; no Hartig net; cauline pericyclic fibers well-developed, fibers associated with leaf bundles; stamens early inverting.

Cassiopoideae + Ericoideae: leaves opposite, revolute [ericoid].

Cassiopoideae Kron & Judd - 1/12. Circumboreal.

Ericoideae Link - 19/1790. Most diverse in South Africa and Malesia to S.E. Asia, also general N. hemisphere to S. South America, inc. Tristan da Cuhna and Falkland Islands.

Harrimanelloideae [Styphelioideae + Vaccinioideae]: K in fruit persisting, not withering.

Harrimanelloideae Kron & Judd - Leaves sublinear; flowers single, axillary. - 1/2. Interruptedly circumboreal.

Styphelioideae + Vaccinioideae: ?

Styphelioideae Sweet - Epidermis lignified; leaf bundles with well-developed abaxial fibrous vascular sheaths, no adaxial cap; leaves xeromorphic, pungent, no clear midrib; A = and opposite sepals, epipetalous, anthers monothecal, appendages 0. - 35/545. Australasia, Pacific, Chile (Lebetanthus).

Vaccinioideae Arnott - Ca 50/1580. N. hemisphere, Malesia and montane Central and South America, Australia (Queensland), few in Africa.

EUASTERIDS: ellagic acid 0, proanthocyanidins not common, caffeic acid +; inflorescence cymose; A epipetalous, = and opposite K or P, [polyandry very uncommon indeed].

EUASTERID I: G [2]; loss of introns 18-23 in d copy of RPB2 gene.

Unplaced (perhaps to be included in Garryales):

Oncothecaceae + Metteniusaceae: nodes 5:5; petiole bundles arcuate, complex; A basifixed, G [5], 2 ovules/carpel; fruit a drupe; endosperm copious.

ONCOTHECACEAE Airy Shaw - Phloem stratified; astrosclereids +; plant glabrous; A extrorse, anthers bisporangiate/monothecal, ovules campylotropous; stone several-seeded. - 1/2. New Caledonia.


METTENIUSACEAE Schnizlein - Stomata anomo-cyclocytic; mesophyll fibers +; hairs ± T-shaped; flowers large, C valvate, contorted, A adnate to C, anthers moniliform, the 4 sporangial series [i.e. vertical rows of locellae] dehiscing individually, 4 carpels very much reduced, barely evident, style long; fruit 1-seeded, asymmetrically ridged; seed coat thin, vascularised; embryo curved. - 1/7. Costa Rica, Panama and N.W. South America.

ICACINACEAE Miers - Endocarp cells papillate. - 23(?24)/149(150). Pantropical, inc. W. Pacific, to China and Japan.

Apodytes group - Fruit very asymmetrical, ribbed, style thin, persistent. - 2/7: Old World tropics, to Australia (Queensland).

Cassinopsis – Stomata cyclocytic; leaves opposite. - 1/4: Africa and Madagascar.

Emmotum groupG [3], 2 ovules/carpel; fruit flattened and symmetrical. - 4(?6)/21(32): Central and South America, West Indies, rarely Malesia.


Woody; group II decarboxylated iridoids [inc. aucubin], gutta +; vessel elements with scalariform perforation plates; nodes?; sclereids +; hairs unicellular; plants dioecious; flowers small; C valvate, free, anthers basifixed, pollen ± atectate, ovary 1-celled, apical crassinucellate apotropous ovules, style at most short; fertilization delayed; fruit indehiscent. - 2 families, 3 genera, 18 species.

GARRYACEAE Lindley - Leaves opposite; G inferior, 1 ovule/carpel. - 2/17. W. North America, Central America and the Greater Antilles (Garrya)and East Asia (Aucuba).

EUCOMMIACEAE Engler - Plant with articulated laticifers; bracteoles 0, P 0; staminate flowers: connective prolonged; carpellate flowers: G [2], one aborts, micropyle long, nucellar cap +, stigmas spreading, decurrent; fruit a samara; seed 1. - 1/1. China.

BORAGINACEAE + VAHLIACEAE + GENTIANALES + LAMIALES + SOLANALES: (8-ring deoxyflavonols +); vessel elements with simple perforations; C tube initiation late [sampling!], [vascularised] disc at base of G, style long.


BORAGINACEAE Jussieu - Isokestose and higher inulin oligosaccharides as storage; sieve elements with nuclear non-dispersive protein bodies; plant often roughly hairy, hairs with a basal cystolith or cystolith-like body, and/or walls calcified; cyme scorpioid, bracteoles 0; K free, anther placentoid 0, pollen with pseudocolpia, 2 epitropous ovules/carpel. - 148/2740. Temperate to tropical.

1. Boraginoideae Arnott - Pyrrolizidine alkaloids, alkannin, prenylated napthoquinones [roots with reddish or purple dye]; corolla mouth with fornices [inpushings of corolla tube], style gynobasic; fruit a schizocarp; endosperm haustoria 0. - 112/1600. Largely (warm) temperate, some on mountains in the tropics.

Goes where? Codonoideae Retief & A. E. van Wyk - K deeply linear-lobed, C 10-12-lobed, A = C, many ovules/carpel; fruits are loculicidal. - 1/2. Southern Africa.

Hydrophyllaceae + Heliotropioideae + Cordioideae + Ehretioideae + Lennoaceae: single layer of transfer cells [cells with labyrinthine ingrowths of the wall] in the testa; endosperm with micropylar and chalazal haustoria.

2. Hydrophyllaceae [sic] R. Brown - G with parietal placentation; fruit a loculicidal capsule. - 17/225. New World, esp. drier areas W. United States.

Heliotropioideae + Cordioideae + Ehretioideae + Lennoaceae: bark oxidises; transfer cells in funicle and placenta also; endocarp multilayered.

3. Heliotropioideae (Schrader) Arnott - Pyrrolizidine alkaloids +; stigma receptive only basi-laterally, discoid, then conical and ± bilobed at sterile apex, or hemispherical, with a ring of hairs; fruit a schizocarp. - 5/405. Tropical to warm temperate.

4. Cordioideae (Link) Chamisso - Terpenoid-based quinones +; fruit drupaceous; cotyledons plicate. - 3/330. Tropical, especially South America.

5. Ehretioideae (Martius) Arnott - Fruit drupaceous. - 8/170. Mostly tropical.

6. Lennooideae L. Craven - Echlorophyllous, herbaceous root parasites; stem with cortical bundles; leaves reduced to scales; inflorescences congested; K long, narrow, free, nectary 0, G [5-16], style stout; K and C persistent in fruit; embryo minute, undifferentiated. - 3/7. S.W. U.S.A. to N. South America.

VAHLIACEAE Dandy - Hairs uniseriate-glandular; flowers small; K valvate, C free, G inferior, 1-locular, placentae apical, ovules bitegmic, micropyle endostomal, endothelium +, styles separate, diverging. - 1/8. Africa and Madagascar to India.


Iridoids, indole alkaloids +; pits vestured; glandular hairs 0; nodes?; leaves opposite, joined by a line across the stem, colleters +; endothelium 0; endosperm nuclear. - 5 families, 1118 genera, 16637 species.

RUBIACEAE Jussieu - Route II carboxylated iridoids +, shikimic-acid derived anthraquinones, isoquinoline alkaloids, etc. +; raphides or crystal sand +; leaves with interpetiolar stipules, innervated from circumferential vascular ring; G inferior. - >600+/10,000+. World-wide, but especially tropical.

1. Rubioideae - Plants Al accumulators; hairs articulated; C valvate, heterostyly esp. common. - Worldwide.

Cinchonoideae Rafinesque

Plants woody; route II carboxylated iridoids, indole alkaloids +; hairs mostly cylindrical; secondary pollen presentation common. - Worldwide.

2. Luculia + Coptasapelta - 2/21. Himalayas, China, to Malesia.

3. Cinchonidinae Robbrecht & Manen - Corynanthean and complex indole alkaloids +; C left contorted. - Pantropical.

4. Ixoridinae Robbrecht & Manen - Pantropical.

Gentianaceae + Loganiaceae + Gelsemiaceae + Apocynaceae: route I secoiridoids +; internal phloem +; C tube formation late, syncarpy postgenital.

GENTIANACEAE Jussieu - Leaves sessile, usu. connate basally; C contorted to the right, marcescent, disc 0; fruit a septicidal capsule; 100 bp deletion in trnL gene. - 87/ca 1650.

1. Saccifolieae (Maguire & Pires) Struwe, Thiv, V. A. Albert & Kadereit - Placentation parietal. - 4/19. tropical South America, Panama.

2. Exaceae Colla - 6/165. Africa, esp. Madagascar, Indo-Malesia, and to Australia and New Zealand (some Sebaea).

Chironieae + Helieae + Potalieae + Gentianeae: xanthones, L-(+)-bornesitol +; placentation parietal.

3. Chironieae (G. Don) Endlicher - Distinctive 6-substituted xanthones. - 23/159. Tropics and warm N. temperate.

Helieae + Potalieae + Gentianeae: nectary +.

4. Helieae Gilg - Style often long, twisted and flattened when dry. - 22/184. Tropical Central and South America, Caribbean.

5. Potalieae Reichenbach - C-glucoflavones +. - 13/154. Pantropical.

6. Gentianeae Colla - Distinctive xanthones, C-glucoflavones +. - 17/950. North temperate, to the Celebes (some Tripterospermum) and Africa and Madagascar (some Swertia).

Loganiaceae + Gelsemiaceae + Apocynaceae: ?

LOGANIACEAE R. Brown - 13/420. Pantropical, esp. Australia and New Caledonia.

Gelsemiaceae + Apocynaceae: nodes 1:1; seeds ± flattened.

GELSEMIACEAE Struwe & V. Albert - Flowers heterostylous, A extrorse or latrorse, style twice bifid. - 3/12. ± tropical; South East Asia, Africa, America, inc. Pteleocarpa.

APOCYNACEAE Jussieu - Steroidal alkaloids, cardenolides, route II decarboxylated iridoids +; K with basal adaxial colleters, C left-contorted, A ± connivent, filament short, pollen transported in foam, apex of G alone postgenitally syncarpous, stylar head swollen, differentiated, stigmatic zone basal; testa multiplicative. - 415/4555. Largely tropical to warm temperate.

1. "Rauvolfioideae" Kosteletzky - 84/980. Tropical.

Apocynoideae + Periplocoideae [Secamonoideae + Asclepiadoideae]: iridoids 0 [this level?]; C right-contorted, anthers with lignified basal appendages [guide rails], adnate to style head, pollen porate, stylar head radially differentiated; fruit a follicle; seeds comose.

2. Apocynoideae Burnett - 77/860. Largely tropical.

3. Periplocoideae R. Brown - Anthers without guide rails, pollen in tetrads, collected on translator [spoonlike structure + basal sticky viscidium]. - 31/180. Old World, tropics to dry temperate.

Secamonoideae + Asclepiadoideae: C tube formation intermediate, A inserted well below bases of corolla lobes, filament 0, staminal feet erect, connate, forming tube around ovary, anthers inserted on top, microsporogenesis successive, pollen inaperturate, psilate, aggregated into pollinia, pollinia of one pollinarium from half anthers of adjacent stamens, orbicules 0, translator [retinaculum] of hardened resinous secretion [mostly stigmatic] and apical adhesive corpusculum, nectaries behind guide rails.

4. Secamonoideae Endlicher - Pollinia 4, lacking outer wall, translator arms [caudicle] 0, pollen in tetrads, granular layer of exine thick. - 9/170. Old World, esp. Madagascar, tropics to temperate.

5. Asclepiadoideae Burnett. - Anthers bisporangiate [1 sporangium from each theca is lost], pollinia 2, caudicle +, anther secreting wall around pollinium, granular layer of exine thin. - 214/2365. Tropics to temperate, drier areas esp. in Africa.

LAMIALES + SOLANALES: iridoids 0; nodes 1:1; K connate, anther sacs with placentoids.

LAMIALES Bromhead  

Cornosides, verbascoside acteoside [caffeic acid ester] +; glandular hairs with vertical divisions in the head +; leaves opposite; endosperm with micropylar haustoria; protein bodies in nuclei; mitochondrial coxII.i3 intron 0. - 22 families, 1059 genera, 23275 species.

PLOCOSPERMATACEAE Hutchinson - Leaves articulated near the base of the petiole; A extrorse, versatile, placentation parietal, 2 ovules/carpel, style divided twice, lobes stigmatic, not expanded; seed with tuft of multicellular hairs at chalazal end. - 1/1. Central America.

[Carlemanniaceae + Oleaceae] [Tetrachondraceae [[Calceolariaceae [Peltanthera + Gesneriaceae]] [Plantaginaceae, Lamiaceae, etc.]]]: cells in heads of glandular hairs with vertical walls only; flowers 4-merous [?: reverses to 5-merous....].

Carlemanniaceae + Oleaceae: A 2, stigma ± clavate; exotestal cells ± palisade.

CARLEMANNIACEAE Airy Shaw - Flowers weakly obliquely monosymmetric; pollen colpate, G inferior. - 2/5. Thailand, Laos, Vietnam, S. China, Sumatra.

OLEACEAE Hoffmannsegg & Link - Flowers 4-merous; anther thecae ± back-to-back. - 24/615. More or less worldwide, especially East Asia.

1. Fontanesieae L. Johnson - Pits vestured; 1 ovule/carpel; fruit a samara. - 1/2. Sicily, W. Asia, China.

2. Forsythieae L. Johnson - 2/8. S.E. Europe, East Asia.

3. Myxopyreae Boerlage - Myxopyroside iridoid pathway +; cortical bundles in corners of angled stem. - 3/7. Indo-Malesia.

4. Jasmineae + Oleeae: oleoside +; 2(-4) ovules/carpel.

5. Jasmineae Lamarck & Candolle - Fruit bilobed; seed coat multilayered, mesotesta with wholly thickened or band-thickened anticlinal walls; 21kb chloroplast inversion. - 1/450. Tropical to warm temperate Old World, some in America.

6. Oleeae (R. Brown) Dumortier - Flavone glycosides +; libriform fibers +; pits vestured; n = 23. - 17/415. Tropical and subtropical, inc. New Zealand and Hawaii.

Tetrachondraceae [Calceolariaceae [Peltanthera + Gesneriaceae] [Plantaginaceae, Lamiaceae, etc.]]: deletion in the matK gene.

TETRACHONDRACEAE Wettstein - Flowers 4-merous, pollen in tetrads, 6-sulcate, psilate, nectary 0, 2 erect ovules/carpel. - 2/3. Patagonia, Australia and New Zealand (Tetrachondra), S. U.S.A. to South America (Polypremum).

HYDROSTACHYACEAE Engler - Submerged herbs; primary root 0; stomata 0; leaves in a rosette, deeply and complexly divided, surface with small enations, stipule +; inflorescence spicate, plants di(mon)oecious, P 0, nectary 0; staminate flowers: A 2, monothecal, extrose, pollen in tetrads, inaperturate; carpellate flowers: G [2], transverse, placentation parietal, many ovules/carpel, styluli separate, filiform; fruit a septicidal capsule; seeds minute, outer cell walls much thickened, mucilaginous; endosperm scanty or 0. - 1/20. C. and S. Africa, Madagascar.

N.B. - this is a possible position....

[[Calceolariaceae [Peltanthera + Gesneriaceae]] [Plantaginaceae [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]]]]: shikimic-acid derived anthraquinones [see also Rubiaceae], 6- and/or 8- hydroxylated flavone glycosides + [? Tetrachondraceae], storage substances stachyose and other oligosaccarides; flowers vertically monosymmetric, 5-merous, C bilabiate, (2:3), adaxial lobes outside the others in bud [ascending cochleate], tube formation late, A 4, didynamous, anthers connivent, many ovules/carpel; endosperm also with chalazal haustoria [?position in tree]; protein bodies in nuclei lamellar.

[Calceolariaceae [Peltanthera + Gesneriaceae]]: cymes pair-flowered.

CALCEOLARIACEAE Olmstead - Lower lip of C saccate, nectary 0, A 2. - 3/200. Upland tropical and W. temperate South America, Brasil, also New Zealand.

[Peltanthera + Gesneriaceae]: suspensor large.

GESNERIACEAE Candolle - Stomata anisocytic; placentation intrusive parietal. - 147/3870. Largely tropical.

Didymocarpoideae + Epithematoideae: 3-desoxyanthocyanins 0, chalcones, aurones +; endosperm inconspicuous, cotyledons unequal.

1. Didymocarpoideae - Ovary wall not richly vascularised, placentae lamelliform-recurved, ovules restricted to distal end, style usually not well set off from ovary; fruit elongated; testa cells ornamented. - 78/1900. Mostly Old World tropics, some New World.

2. Epithematoideae - Dihydroxyphenolics [e.g. acteoside] 0; secretory canals, medullary bundles +; ovary short, abruptly narrowed into the style. - 6/75. India, South East Asia to Malesia, 1 sp. W. Africa, 1 Central America to Peru.

3. Gesnerioideae Link - 3-desoxyanthocyanins +, chalcones, aurones 0; seeds without surface ornamentation; endosperm conspicuous; GCyc2 gene lost. - 63/1250. Predominantly Neotropical, a few S.W. Pacific, East Asia.

3a. Coronathereae - Trees or ± shrubby; stomata anomocytic; nectary embedded in G wall, vascularized from A traces; capsules septicidal (and loculicidal; fruit a berry); n = 37(-45). - 9/20. Solomon Islands, Antilles, New Caledonia, S. South America.

Titanotricheae + Gesnerieae: (plant with scaly rhizomes).

3b. Titanotricheae - Inflorescence racemose, with bulbils; testa striate-reticulate; n = 20. - 1/1. China, Japan, Taiwan, scattered in W. Malesia.

3c. Gesnerieae - G superior to inferior, numerous vascular bundles in ovary wall from which nectary is vascularized; n = (8) 9 (10) 11 (12) 13-14 (16). - 53/1500. New World.

Peltanthera - 1/1. Central and W. South America.

[Plantaginaceae [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]]]]: route II decarboxylated iridoids, 6- or 8-hydroxyflavones or 6 methoxyflavones +, cornosides generally 0.

[Plantaginaceae [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae]]]]]: ?

PLANTAGINACEAE Jussieu - Hairs with gland head not vertically divided. - Ca 47/1350. Mostly temperate.

[Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae]]]]: ?

SCROPHULARIACEAE de Jussieu - 65/1700. World wide.

[Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae]]]: ?

STILBACEAE Kunth - 11/39. Most South Africa, esp. the Cape Province, to tropical Africa, Madagascar, the Mascarenes, and Arabia.

[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]]: ?

LAMIACEAE Martynov - Pollen colpate, G with two ovules/carpel. - 236/7173. World-wide.

1. Symphorematoideae Briquet - Lianes; inflorescences in 3-7-flowered capitate cymes with an involucre of bracts; flowers polysymmetrical, K 5-8, C 5-16, A 4-18, disc 0, G imperfectly 2-locular, ovules apical, atropous. - 3/27. India, Sri Lanka, South East Asia, Malesia.

2. Viticoideae Briquet - 10/376-526. Tropical, esp. South East Asia-Australia.

3. Ajugoideae Kostel. - Exine with branched columellae. - 24/1115. Cosmopolitan, but many temperate, and esp. South East Asia to Australia.

4. Prostantheroideae Luersson - Flowers polysymmetric. - 16/317. Australia.

5. Scutellarioideae (Dumortier) Caruel - K two-lipped; seeds tuberculate. - 5/380. ± Cosmopolitan.

6. Lamioideae Harley - Laballenic acid +; style gynobasic. - 63/1210. Esp. Europe to Asia, some cosmopolitan, but v. few Antipodean.

7. Nepetoideae (Dumortier) Luersson - Commonly aromatic [volatile terpenoids, rosmarinic acid], iridoid glycosides, acteosides 0 (+); stem endodermis +; pollen trinucleate, hexacolpate, style gynobasic; exocarp with mucilaginous cells producing hygroscopic spiral fibrils; cotyledons investing embryo. - 105/3675. World-wide, but esp. (warm) temperate.

[Paulowniaceae, Phrymaceae, Orobanchaceae]: ?

PAULOWNIACEAE Nakai - 1/6. (Warm) temperate East Asia.

PHRYMACEAE Schauer - Ca 19/234. ± World-wide, esp. temperate and W. North America and Australia, but few humid tropics.

OROBANCHACEAE Ventenat - Stomata permanently open; orobanchin +; glandular hairs lacking vertical walls in the heads; inflorescence racemose; abaxial lobes of C often outside others in bud [aestivation descending cochleate], anthers with basal tails/awns, pollen colpate. - 99/2061. World wide, but especially N. (warm) temperate and Africa-Madagascar.

[Thomandersiaceae + Verbenaceae]: ?

THOMANDERSIACEAE Sreemadhavan - Woody; phloem stratified; stomata anisocytic; lamina with flat glands abaxially, petiole swollen apically and basally; inflorescence racemose; K with nectaries on the outside, pollen 5-6-colpate, 1-3 hemianatropous ovules/carpel; fruit with "jaculators" [cup-shaped expansion of funicle], K accrescent; seed with rather large hilum; seed coat with ascending-imbricate scales or warts, exotesta palisade, not lignified, up to 6 layers of cells in the warts; embryo strongly curved, cotyledons thin-foliaceous, complexly folded. - 1/6. W. and C. Africa (Map: from Heine 1966).

VERBENACEAE Jaume Saint-Hilaire - Non-glandular hairs unicellular; inflorescence racemose; pollen exine thickened near apertures, G with two ovules/carpel, stigmatic head swollen; fruit a schizocarp/drupe; endosperm 0. - 30/1100. Pantropical (to warm temperate), but mostly New World.

LINDERNIACEAE Borsch, K. Müller, & Eb. Fischer - Iridoids 0; inflorescence racemose or flowers from the axils of leaves; glandular hairs on the inside of the C, embryo sac spathulate, stigma bilobed, sensitive; seeds with ruminate endosperm. - ca. 6/170. Pantropical to warm temperate.

BIGNONIACEAE Jussieu - Leaves bicompound; tapetum amoeboid, pollen colpate, bundles in the ovary wall and also opposite septum, 2 series of ovules/carpel; endosperm 0; cotyledons obcordate, lobed; n = 20. - 110/800. Mainly tropical, esp. South America.

1. Jacarandeae Fenzl - Staminode large, bearded; n = 18. - 1/50. Tropical America, esp. South America.

2. Tourretieae K. Schumann - Tendrillar vines; inflorescence a bracteate raceme. - 2/6. Andes in South America and N. to Mexico.

3. Bignonieae Dumortier - Tendrillar lianes; anomalous secondary thickening + [basal condition is for the xylem cylinder to be 4-lobed]; leaves usu. ternate; fruit usu. septicidal, with persistent septum and whip-like strands [= vascular bundles opposite septum]. - 21/360. American, largely tropical.

4. The rest - Pantropical.

SCHLEGELIACEAE Reveal - Woody; nectaries on outside of K; fruit baccate. - 4/25. Mexico to tropical South America, Cuba (Synapsis).

ACANTHACEAE Jussieu - Acicular fibers +; stomata diacytic; capsule dehiscence explosive; endosperm development asymmetrical [the two haustoria come to lie close to each other]. - 229/3500. Mostly tropical.

1. Nelsonioideae Sreemadhavan - Inflorescence bracts spiral; adaxial lobes of C outside others in bud [aestivation descending cochleate]. - 6-7/170. Tropical.

Acanthoideae[Thunbergioideae + "Avicennioideae"]: pollen other than colporate, 2 collateral ovules/carpel, endothelium 0; endosperm 0.

2. Acanthoideae - Capsules obovoid, seeds borne on hook-like hardened funicles. - 217/3220. World-wide; the bulk of the family.

2A. Acantheae - Nodes not swollen; anthers monothecous, pollen tricolpate. - 21/500.

[Ruellieae + Justicieae] [BAWN clade]: cystoliths; pollen porate.

Ruellieae + Justicieae: ?

2B. Ruellieae - Filament curtain +, pollen often reticulate, (ovules several); seeds with hygroscopic trichomes. .

2C. Justicieae - Anther thecae not equal in height, pollen tricolporate, hexapseudocolpate.

BAWN clade: ?

2D.Nemacanthus - K united 3 + 2, pollen tricolporate, intercolpal regions psilate/foveolate; seed with hygroscopic trichomes. - 1/30. Africa, Madagascar, Arabia to Vietnam.

2E. Whitfieldieae - Pollen biporate, lenticular, granular around apertures; seeds with concentric rings of ridges.

Andrographidae + Barlerieae: ?

2F. Andrographidae - Pollen colporate, ornamented and thickened exine surrounding or over apertures; 3+ ovules/carpel. - /75. Asia.

2G. Barlerieae - C quincuncial: seed with hygroscopic trichomes. - /420. Pantropical.

Thunbergioideae + Avicennioideae: cotyledons folded.

3. Thunbergioideae Kostel. - Vines; inflorescence with 2 or more flowers in the median plane of the leaf axil, adaxial flowers opening first; bracts 0, bracteoles large, connate, K a rim, anthers with lignified unicellular hairs, sagittate, (elongated) porose, endothecium 0. - Ca 5/150. Tropical America, Africa and Madagascar, fewer in South East Asia - Malesia.

4. Avicennioideae - Wood with successive cambia, phloem islands in bands of conjunctive tissue; leaves with salt glands on both sides; A = and alternating with C, ovule lacking micropyle; K persistent, green; seed 1. - 1/8. Mangroves in tropics, but also warm temperate.

PEDALIACEAE R. Brown - Hairs mucilaginous; nectary glands at base of pedicel; connective with apical gland, pollen 5-13-zonocolpate. - 13/75. Mostly tropical, in coastal or arid habitats, Old World.

MARTYNIACEAE Horaninow - Inflorescence racemose; pollen inaperturate, exine dissected into 20-40 platelets, placentation parietal; fruit with apical hooks, mesocarp peeling off, endocarp woody. - 5/16. Tropical and subtropical America.

BYBLIDACEAE Domin - Rhizomatous herbs; stomata paracytic; leaves linear, abaxially curved, veins parallel; flowers single, axillary, polysymmetric, bracteoles 0; C contorted, connate only basally, A 5, nectary 0. - 1/6. Australia, southern New Guinea.

LENTIBULARIACEAE Richard - Insectivorous rosette herbs, primary root not developing; inflorescence racemose; C with an abaxial nectar-secreting spur, A 2, disc 0. - 3/320. World-wide.

SOLANALES Dumortier  

O-methyl flavonols +; pollen tube with callose; K persistent in fruit; endosperm development? - 5 families, 165 genera, 4080 species.

Montiniaceae + Sphenocleaceae + Hydroleaceae: petiole bundle(s) arcuate; inflorescence terminal.

MONTINIACEAE Nakai - Flowers imperfect; C valvate, A free, extrorse, G inferior, style short, stout; seeds winged. - 3/5. Africa and Madagascar.

Sphenocleaceae + Hydroleaceae: placentae swollen [?level]; endosperm cellular, at most scanty, with multicellular micropylar and chalazal haustoria.

SPHENOCLEACEAE Baskerville - Rather fleshy annual herbs; fructose +; stomata tetracytic; inflorescences spicate; C free, A free, nectary 0, G ± inferior, style short; fruit capsular, circumscissile. - 1/2. Pantropical.

HYDROLEACEAE Berchtold & J. Presl - Filaments swollen at the base, stylodia +. - 1/12. Tropical.

Convolvulaceae + Solanaceae: caffeic acid esters, tropane [polyhydroxynortropanes] and pyrrolidine alkaloids, acylated anthocyanins +; internal phloem +; C usu. contorted or induplicate-valvate; K notably conspicuous in fruit; young seeds starchy, (endosperm development notably variable).

CONVOLVULACEAE Jussieu - Plant laticiferous; K quincuncial, large, placentoid 0; chloroplast gene rpl2 intron absent. - 57/1600. World wide.

1. Humbertioideae Roberty - Large tree; vascular bundles collateral; petiole bundle annular; latex in the flowers alone; flowers single, axillary, strongly obliquely monosymmetric, A adnate to base of C, filaments bent in bud, style clavate; fruit a few-seeded drupe. - 1/1. Madagascar.

2. Convolvuloideae - Herbaceous (woody) and right-twining vines; pollen pantoporate or 6-polycolpate, (1-)2(-4) erect crassinucellate ovules/carpel. - 56/1600. World-wide.

SOLANACEAE Jussieu - 102/2460. World-wide, but overwhelmingly tropical America.

1. Schwenckioideae - 4/31. South America.

Schizanthoideae + Goetzeoideae + Cestroideae + Petunioideae + Solanoideae + Nicotianoideae: ?

2. Schizanthoideae (Miers) Hunziker - Flowers resupinate, strongly monosymmetric, abaxial pair of C connate, forming a keel, A 2 [abaxial-lateral], staminodes 3. - 1/12. Chile.

3. Goetzeoideae (Airy Shaw) Thorne & Reveal - 4/7. Greater Antilles, but not Jamaica, South America.

4. Browallioideae Kosteletzky - 8/195. South and Central (and North) America.

5. Petunioideae - 13/160. Central and South America.

Solanoideae + Nicotianoideae: x = 12.

6. Nicotianoideae Miers - 8/125. Mostly Australian, also North and South America, Africa.

6. Solanoideae Kosteletzky - Leaves often geminate and/or branching apparently not axillary; fruit a berry; seeds flattened; embryo curved. - 61/1925. World-wide, but esp. South America and others N. temperate.

EUASTERID II: vessel elements with scalariform perforations; flowers rather small, style short; endosperm copious, embryo short/very short.


Iridoids?; petiole bundles arcuate; C valvate, free, ?development, A free from P/C, 1-2 apical apotropous ovules/carpel; fruit a drupe, stones one-seeded. - 5 families, 21 genera, 536 species.

Cardiopteridaceae + Stemonuraceae: Nodes 3:3; stomata cyclocytic to anisocytic; hairs unicellular; leaf margins entire; apex of petals inflexed, A basifixed, G [?2], unilocular, with 2 apical crassinucellate ovules, funicular obturator +; fruit a 1-seeded drupe; testa vascularised.

CARDIOPTERIDACEAE Blume - 5(?6)/43(44). Tropics, inc. the Pacific, to Taiwan.

STEMONURACEAE Kårehed - Style 0, stigma broad. - 12/80. Tropics, esp. Indo-Malesia to Australia (Queensland).

Aquifoliaceae [Phyllonomaceae + Helwingiaceae]: nodes 1:1; leaf margins toothed, stipules small, cauline; drupe with separate pyrenes.

AQUIFOLIACEAE A. Richard - Pollen conspicuously gemmate/clavate. - 1/405. ± World-wide, esp. America and South East Asia-Malesia, few in Africa.

Phyllonomaceae + Helwingiaceae: hairs 0; lamina with brochidodromous venation, stipules fimbriate; inflorescence epiphyllous, on adaxial side of lamina.

PHYLLONOMACEAE Small - Petiole bundle annular; G inferior, several ovules/loculus, stigmas recurved; fruit a berry; testa multilayered, exotestal cells large, thick-walled, mucilaginous, 2-3 layers of flattened cells; endosperm hemicellulosic. - 1/4. Mexico to Peru.

HELLWINGIACEAE Decaisne - P 3-5, A = and alternating with P, pollen with diffuse endoapertures. - 1/3. Himalayas to Japan.

ASTERALES [ESCALLONIACEAE [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]: iridoids +; C tube initiation early, G [2-3], inferior.


Route I secoiridoids, oligo- or polyfructosans, inc. inulin, with isokestose linkages +; nodes 3:3; flower size?, C valvate, petal apiculi inflexed, A free from C, pollen grains trinucleate, disc +, many ovules/carpel, integument <7 cells thick, endothelium +, antipodal cells ephemeral, style long; endosperm copious, micropylar and chalazal haustoria +; mitochondrial rpl2 gene lost. - 11 families, 1649 genera, 25790 species.

Rousseaceae + Campanulaceae: flowers large, A free.

ROUSSEACEAE Candolle - 4/13. Mauritius, New Guinea to New Zealand.

1. Rousseaoideae - Anthers basifixed, attached their entire length to stout connective, sagittate, extrorse, pollen 6- or 8-porate. - 1/1. Mauritius.

2. Carpodetoideae (Fenzl) J. Lundberg - Flowers small. - 3/12. New Zealand, E. Australia, Papuasia.

CAMPANULACEAE Jussieu - Polyacetylenes +, iridoids 0; articulated laticifers +; crystals acicular; flowers monosymmetric, median K abaxial, secondary pollen presentation + [A basifixed, hairs at the tip of the style, anthers at least initially close to stigma], G [2]; expansion of chloroplast inverted repeat into small single copy region, 5bp ndhF deletion, chloroplast accD gene lost, infA gene defunct, biparental plastid transmission [?levels]. - 84/2380. World-wide.

Nemocladoideae + Campanuloideae: pollen spheroid to oblate-spheroid, with spicules, style with long hairy part, hairs retractile [inc. Nemacladoideae?]; fibrillar protein bodies in nuclei.

1. Nemacladoideae M. H. G. Gustafsson - Flowers small, C 3:2, filaments forming tube, anthers flip back after pollen release, style head bending towards or away from median. - 3/15. S.W. U. S. A., Mexico.

Nemacladus has groups of remarkable reflexed pseudonectaries at the bases of two filaments.

2. Campanuloideae Burnett - Flowers polysymmetric, median K adaxial, stamens often sprawling at bottom of corolla tube after pollen is shed although bases conceal nectar, G [3-5]; extensive rearrangements in the chloroplast inverted repeat. - 50/1050. Especially N. temperate, very few in the Australia-New Zealand area.

Lobelioideae + Cyphocarpoideae + Cyphioideae: pollen prolate.

3. Lobelioideae Schönland - Chelidonic acid, pyrrolizidine alkaloids +; flowers resupinate, anthers connate, style with brush hairs and pollen in pollen box. - 329/1200. Almost world-wide, but largely tropical.

4. Cyphocarpoideae Miers - Leaves deeply lobed; C 1:4, adaxial petal cucullate, with apical appendage, A epipetalous, free, ovary notably elongated; fruit dehiscing through sides; nuclear inclusions fibrillar. - 1/3. Chile.

5. Cyphioideae Schönland - Roots tuberous; A basally connate, pollen psilate, style bends away from median K but does not elongate after A dehiscence, stigma with a fluid-filled terminal cavity with a lateral pore; fruit a septi- and loculicidal capsule [valves bifid]. - 1/65. Africa, esp. South Africa.

PENTAPHRAGMATACEAE J. Agardh - Rather fleshy herbs, rooting at base of stem; wood rayless; leaves asymmetrical, 2-ranked; inflorescences cymose, scorpioid; K petaloid, 2 large + 3 small, nectariferous cavities between septae joining hypanthium to G, A adnate to C, extrorse, basifixed, embryo sac protruding from micropyle; unicellular micropylar haustorium only, endosperm starchy. - 1/30. South East Asia to Malesia, esp. W. Malesia.

Alseuosmiaceae + Phellinaceae + Argophyllaceae + Stylidiaceae + Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae: ?

Alseuosmiaceae + Phellinaceae + Argophyllaceae: hairs uniseriate, egalndular; lamina serrate, gland-toothed; x = 8.

ALSEUOSMIACEAE Airy Shaw - Endodermis in both stem and leaf; leaves with axillary tufts of uniseriate hairs; C lobe margin ± erose, winged, A adnate to C. - ?5/10. New Guinea, E. Australia, New Zealand, New Caledonia.

Phellinaceae + Argophyllaceae: pollen (spiny) with rugulose exine, ovules apotropous, style short.

PHELLINACEAE Takhtajan - Benzylisoquinoline [homoerythrina] alkaloids +; guard cells huge[!]; plant dioecious. - 1/12. New Caledonia.

ARGOPHYLLACEAE Takhtajan - Hairs T-shaped, multicellular, with slits over the stalk cell. - 2/17. S.W. Pacific.

STYLIDIACEAE R. Brown - A extrorse, fewer than C [2-3], pollen colpate. - 6/245. South East Asia to New Zealand, S. South America.

1. Donatioideae B. Chandler - Flowers solitary, terminal; sepals and petals free; suspensor short. -1/2. New Zealand, Tasmania, S. South America.

2. Stylidioideae - Flowers semi-resupinate, split-monosymmetric, A 2, adnate to style, anther thecae set end to end, pollen 2-8-colpate. - 5/155. Esp. Australia, also South East Asia, Malesia, S. South America.

Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae: caffeic acid +; vessel elements with simple perforations; C connate, with strong ± fused marginal [commissural] veins joining the median near the apex, A adnate to C, tapetum multinucleate, pollen spiny, G [2], integument >10 cells thick, vascular bundle proceeding to the micropyle; no endosperm haustoria, embryo long.

MENYANTHACEAE Berchtold & J. Presl - Aquatic herbs; leaf bases broad; flowers heterostylous, corolla lobes with marginal wings or fringes, G superior, placentation parietal. - 5/ca 40. World-wide.

Goodeniaceae + Calyceraceae + Asteraceae: secondary pollen presentation + [details vary considerably], pollen with bifurcating columellae, stigma dry, ± papillate; K persistent in fruit; x = 8.

GOODENIACEAE R. Brown - Corolla slit-monosymmetric [0:5], lobes with marginal wings or fringes, stylar cup +. - 12/440. Largely Australian, Scaevola pantropical.

Calyceraceae + Asteraceae: flowers small, C tubular, commissural veins connate (median veins 0), filament collar +, G with only 1 ovule; fruit a cypsela, K persistent, modified, involved in dispersal.

CALYCERACEAE Richard - Inflorescence an involucrate capitulum made up of cymose units; K connate, aerenchymatous or spine-like, C tubular, outer layer separates and photosynthesises, filaments connate, glands internally alternating with filaments, pollen with intercolpar depressions, ovule apical, apotropous; apex of fruit with a conical body [persistent base of C and style]. - 6/60. South America.

ASTERACEAE Martynov - Sesquiterpene lactones, terpenoid essential oils, iridoids 0; inflorescence capitulate, involucrate, ebracteate; K reduced, anthers connate, with conspicuous apical and basal [calcarate] appendages, caudate, endothecial cells elongated parallel to main axis of anthers, tapetum amoeboid, ovule basal, epitropous, inner surface of style branches stigmatic; fruit with a pappus of bristles. - 1620/22750. World-wide.

1. Barnadesioideae (Bentham and J. D. Hooker) Bremer & Jansen - Long tricellular hairs [epidermal cell is undistinguished, basal cell is short and thick-walled, the other cell is longer and with thin walls: esp. on flowers, even on the bristles of the cypsela]; pollen with intercolpar depressions; cypsela with spines. - 9/94. South America, esp. Andean.

"The Stiftia group" + Mutisioideae + Gochnatioideae + Hecastocleioideae + Carduoideae + Pertyoideae + Gymnarrhenoideae + Cichorioideae + Corymboideae + Asteroideae: style with sweeping hairs, style branches often long; cypsela with twin hairs, pappus capillary (scales, etc.); chloroplast DNA inversion +.

2. "The Stiftia group" - South America, Asia.

3. Mutisioideae - 58/750. - South America.

NOTE: some genera need to be placed in the preceding group.

3. Gochnatioideae (Bentham & J. D. Hooker) Panero & Funk - Style apex rounded; pappus of bristles. - 4(?5)/90. Central and South America, esp. the Caribbean and southern South America.

Hecastocleioideae [Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymboideae + Asteroideae]]]]]: deletion in rpoB gene.

4. Hecastocleioideae Panero & Funk - Style branches rounded; pappus of scales. - 1/1. S.W. U.S.A.

Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymboideae + Asteroideae]]]]: sweeping hairs +; deletion and insertion in rpoB gene.

5. Carduoideae Kitamura - 83/2500. World-wide, but most N. hemisphere, esp. Eurasia/N. Africa.

Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymboideae + Asteroideae]]]: ?

7. Pertyoideae Panero & Funk - Flowers not bilabiate, corolla deeply but unequally divided; pappus of (plumose) bristles. - 5-6/70. Afghanistan to East (and Southeast) Asia.

Gymnarrhenoideae [Cichorioideae [Corymboideae + Asteroideae]]: ?

8. Gymnarrhenoideae Panero & Funk - Plant amphicarpic; heads grouped into synflorescence; pappus of bristles. - 1/1. North Africa to the Middle East.

Cichorioideae [Corymboideae + Asteroideae]: deletion in ndhF gene.

9. Asteroideae - Pollen with cavities separating columellae from foot layer [caveate], with a double tectum, style hairs often rounded, only ± at style tip, stigmatic areas in two marginal bands; rbcL 6bp x 4 inversion. - 1135/16200. World-wide.


ESCALLONIACEAE Dumortier - Cork superficial; nodes 1:1; petals free. - 9/130. Réunion, E. Himalayas and S. China to N.E. Australia and New Zealand, Central and South America.



Nodes 1:1; anthers basifixed.

BRUNIACEAE Berchtold & J. Presl - Stomata cyclocytic, orientation transverse; leaf apex with localized cork cambium; ovules crassinucellate. - 12/75. South Africa, almost entirely the Cape Province.

COLUMELLIACEAE D. Don - Cork pericyclic; nodes 1:1; pericyclic fibers 0; petiole bundles arcuate; bud without scales; leaves opposite, margins toothed, bases meeting; inflorescence terminal, cymose; flowers medium-sized, anther connective well-developed, style long, many ovules/carpel; K persistent in fruit; seeds many, anticlinal walls of exotestal cells thickened, with plasmodesmata. - 3-5. Costa Rica to Chile, in the Andes.



Group II decarboxylated iridoids +; ?nodes; ?stomata; inflorescence terminal, paniculate; plants dioecious; pedicels articulated; flowers small, K small, C apparently free [one position], A free from C, G [3], abaxial carpel fertile, 1-2 apical apotropous ovules/carpel, nucellus type?, funicular obturator +; fruit a single-seeded drupe; endosperm nuclear; mitochondrial rpl2 gene lost. - 7 families, 494 genera, 5489 species.

PENNANTIACEAE J. Agardh - Ovary superior. - 1/4. E. Australia, New Zealand.

Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]]]: young stems with peripheral collechyma; pericyclic fibers 0 or few; nodes 5(+):5(+); leaf bases broad; C apparently free [another position], G inferior, styles/stigmas recurved.

TORRICELLIACEAE H. H. Hu - 3/10. Madagascar, South East Asia and W. Malesia.

Griseliniaceae [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]]: petroselenic acid [in seed] +; C 1-veined, nectary of sorts +, endothelium +.

GRISELINIACEAE A. Cunningham - Stomata encyclocytic; hairs unicellular; leaves 2-ranked, conduplicate; ovule crassinucellate. - 1/6. New Zealand and S. South America.

Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]: ethereal oils, polyacetylenes, acetate-derived anthroquinones, furanocoumarins +, iridoids, tannins 0; lateral roots originating from either side of the xylem poles; schizogenous secretory canals in pericycle; leaf margins toothed or otherwise incised; ultimate units of inflorescence umbels, flowers perfect and imperfect [plant andromonoecious or dioecious]; G [2], both fertile, with nectariferous stylopodium, 2 epitropous ovules/carpel; fruits ± laterally compressed; hemicellulosic seed reserves common; x = 12.

PITTOSPORACEAE R. Brown - Leaf base narrow, margin entire, venation pinnate; inflorescence units not umbellate; flowers medium-sized, K large, G superior, flank nectary +, placentation parietal, many ovules/carpel, endothelium 0, style +; fruit a capsule; seed with fleshy coat. - 9/200. Old World, especially Australia, tropical and warm temperate, outside the immediate Australian region, mainly Pittosporum.

Araliaceae [Myodocarpaceae + Apiaceae]: polyacetylene C18 tariric acid +; leaves (compound), margins toothed or otherwise incised; ultimate units of inflorescences umbels; nectariferous stylopodium +, two epitropous ovules/carpel, one much reduced; fruits ± laterally compressed; 92 bp deletion in rpl16 gene.

ARALIACEAE Jussieu - C valvate. - 43/1450. Largely tropical, few temperate.

1. Hydrocotyloideae Link - Herbs. - 2/175. Tropical, including montane, warm temperate.

2. Aralioideae Eaton - 41/1275. Largely tropical, few temperate.

Myodocarpaceae + Apiaceae: ?

MYODOCARPACEAE Doweld - 3/19. New Caledonia, E. Malesia, and Queensland, Australia.

APIACEAE Lindley - Umbels often compound; C clawed, with inflexed tips, valvate, A inflexed in bud; fruit dry, schizocarpic. - 434/3780. World-wide, esp. N. temperate.

1. Mackinlayoideae Plunkett & Lowry - 5/67. Most S. Pacific Rim, Centella esp. S. Africa, C. asiatica pantropical.

Azorelloideae + Saniculoideae + Apioideae: fruits dorsally flattened.

2. Azorelloideae Plunkett & Lowry - Ca 17/155. South American-Australian, Antarctic islands; Drusa glandulosa from the Canary Islands and Somalia!

Saniculoideae + Apioideae: stipules 0; fruits with secondary [i.e. lateral] ribs +, endocarp of single cell layer, calcium oxalate as druses in pericarp and around commissure, intrajugal vittae [oil ducts in the ribs] +.

3. Saniculoideae Burnett - Inflorescence apparently a simple umbel [probably reduced compound umbels], with showy inflorescence bracts; style separated from disc by a narrow groove. - 11/335. World-wide.

3a. Steganotaenieae C. I. Calviño & S. R. Downie - Plant woody; fruits 2-3 winged [wings exo- and mesocarp alone], intrajugal secretory ducts/cavities associated with the wings much expanded, carpophore +. 2/2-3. Tropical Africa, S.W. Cape.

3b. Saniculeae Burnett - (R)-3'-O-ß-D-glucopranosylrosmarinic acid +; leaf blades often broad, with hairy or spiny tips to the teeth; carpellate flowers sessile; fruit scaly or spiny, carpophore 0, endocarp not lignified. - 8/333. World-wide.

4. Apioideae Seemann - Methylated flavonoids, furanocoumarins, phenylpropenes +; umbels compound; hypostase +, stylopodium lacking groove; carpophore free, bifid [mericarps attached at apex]; x = 11. - 400/3200. Worldwide, esp. N. temperate.

[PARACRYPHIALES + DIPSACALES]: nodes 3:3; true tracheids +; leaves serrate; inflorescence terminal.


Inflorescence racemose; flowers 4-merous, filaments stout, thecae ± embedded in connective; capsule septicidal. 1 family, 3 genera, 36 species.

PARACRYPHIACEAE Airy-Shaw - Leaves spiral. - 3/36. S.W. Pacific.


Group I secoiridoids +; nodes 3:3; buds with scales; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; fruit indehiscent, K persistent; testa vascularised, exotestal cells enlarged, variously thickened and lignified. - 2 (7) families, 45 genera, 1090 species.

ADOXACEAE Trautvetter - Sepals with 1 vascular trace, C rotate, ovary half inferior, style short; chromosomes continuously condensed from prophase to metaphase, etc. - 5/200. North temperate, tropical montane, but not in Africa.

1. Viburnum - Hairs stellate and variants; 1 carpel fertile. - 1/175. North temperate, tropical montane, but not in Africa.

2. Adoxa, etc. - Leaves compound; A extrorse, G [5], embryo sac tetrasporic, but eight nucleate. - 4/23. Mainly north temperate, esp. China, also subtropical.

Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Valerianaceae + Dipsacaceae]]]] [= Caprifoliaceae s.l.]: bark papery-flaky; O-methylated flavones, flavonols +; cork deep-seated; flowers rather weakly monosymmetric, medium-sized, C tubular, nectary as unicellular hairs, anthers sagittate, tapetum amoeboid, pollen grains spheroid, echinate, G inferior, style long; chromosomes "small"; horizontal transfer of mitochondrial rps11 gene [?Morinaceae and upwards in the tree].

DIERVILLACEAE (Rafinesque) Pyck - Fruit septicidal down sides. - 1-2/16. East Asia, S.E. U.S.A.

Caprifoliaceae [Linnaeaceae [Morinaceae [Valerianaceae + Dipsacaceae]]]: ?

CAPRIFOLIACEAE Jussieu - 5/220. Mostly N. temperate, esp. East Asia and E. North America.

Linnaeaceae [Morinaceae [Valerianaceae + Dipsacaceae]]: supernumerary bracts from aborted flowers +; A 4, didynamous, G [3], 2 abort; fruit a cypsela.

LINNAEACEAE (Rafinesque) Backlund - Nectariferous petal abaxial. - 4-5/36. Temperate South East Asia (N. Temperate, inc. Mexico).

Morinaceae + Dipsacaceae + Valerianaceae: perennial rosette herbs with taproot; monoterpenoids, cathecolic tannins, alkaloids +; flank-bridge in stem between lateral bundles; bud scales 0; filaments glabrous, anthers dorsifixed, nectary around base of style, G with sterile loculi much reduced, 1 apical ovule/flower, prominent endothelium with crystal layer, chalazal nuclei variously dividing; exotesta not thickened; endosperm scanty, embryo large; distinctive expansion of the chloroplast inverted repeat.

MORINACEAE Rafinesque - Ethereal oils +, iridoids 0; epicalyx 12-ribbed, calyx monosymmetrical, extra thickened layer of pollen wall, tapetum ?glandular, pollen porate, structures like pollen tubes produced before pollen germination, chalazal cells producing multicellular structures. - 2/13. Balkans to China.

Dipsacaceae + Valerianaceae: saccharose +, quercetin 0; flowers rather small, A ± equal in length, exine columellae prominent and branched, chalazal cells 2-4-nucleate; embryo green.

DIPSACACEAE Jussieu - Starch almost 0; inflorescence capitate, epicalyx 8-ribbed; lateral abaxial C lobes overlapping adaxial lobes, micropyle very long, chalazal nuclei polyploid; endosperm +. - 11/290. Eurasia, Africa, esp. Mediterranean region.

VALERIANACEAE Batsch - Foetid monoterpenoids and sesquiterpenoids, valpotriates +; bracteoles +, epicalyx 0; C spurred, pollen colpate. - 17/315. N. temperate, Andean South America.


MEDUSANDRACEAE Brenan - Secretory canals +; K open, A = and opposite C, dehiscing by recurved valves, staminodes very long, opposite K, densely hairy, disc 0. - 1/2. Tropical W. Africa.

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