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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SUMMARY OF APOMORPHIES IN /APweb/

Note that there may be more detail in the characterisations, even those of the orders, on the individual order pages than appears below as possible apomorphies. This is because the ordinal pages are much more works in progress when it comes to character abstractions. These pages should always be consulted for other characters that may be apomorphies at various levels.

When scanning the summary of potential apomorphies below, remember the sometimes extreme problems one currently faces when trying to place characters on the tree. I repeat some of the points I made in the Introduction:

1. I may not have found all the information about a particular character, there may be disagreement over its interpretation (e.g. Austrobaileyales), or I have added unpublished information (e.g. Myrsinaceae and their relatives, nodal anatomy, etc.; Diapensiaceae, leaf ptyxis; Peridiscaceae, Centroplacaceae, etc.).

2. The sampling of nearly all molecular studies is incomplete (see Salisbury & Kim 2001 for problems caused by sampling). But not only is the sampling in molecular studies often less than we might wish, that of the morphological and chemical characters whose evolution we are interested in understanding is often also very poor.

3. I am fitting characters to a very conservative tree with many polychotomies, although the nodes that are utilised are for the most part strongly supported. This makes optimisation of characters, the assigment of character state change to a particular node on the tree, notably difficult (e.g. Madison & Madison 2002). In nearly all studies of the evolution of characters, the distributions of those characters are optimised on a more or less fully resolved tree, and the construction of supertrees may yield much more detailed hypotheses of relationships. Of course, some nodes on such fully resolved trees and/or supertrees may have little support, and optimisations based on such trees may carry correspondingly little conviction... To see the kind of thing that may happen as our knowledge of relationships improves, note the difference below between the possible synapomorphies given on the Dipsacales page for Dipsacales, and those that might remain if [Columelliaceae + Desfontaineaceae] were sister to Dipsacales. Even in some trees included here, parts have poor support, e.g. Ceratophyllaceae as sister to monocots, relationships within aquatic Alismatales, etc.

4. Exactly how one goes about optimising a character on a tree is critically important. Thus using either parsimony or maximum likelihood, making apparently reasonable suggestions about weighting gains over losses (or vice versa), or more complex assumptions along similar lines, or simply using ACCTRAN or DELTRAN to place the character on the tree (e.g. Donoghue & Ackerley 1996; Cunningham et al. 1998; Omland 1997, 1999; Ree & Donoghue 1999; Polly 2001; Webster & Purvis 2001; Ronquist 2004; Crisp & Cook 2005), may greatly affect the position of synapomorphies on trees, and hence our ideas of evolution. I have tried to indicate where there are particularly important uncertainties as to where characters should be placed on the tree.

5. Finally, although I have paid quite a lot of attention to the delimitation of the character "states" that make up all characterizations, I have not spent enough time on this critical operation. If we are interested in understanding evolution, then fitting the basic variation - not character states - to a tree in principle allows greater flexibility in understanding morphology in the context of local phylogenies (see also Stevens 2000; Endress 2005c). However, many states used here still owe their circumscriptions if not to their delimitation in the context of global phylogenetic analyses, at least to thinking about the variation shown by a particular character across all angiosperms. Such states often have arbitrary limits, and serve best to communicate "information"; whether they are in fact suitable for either phylogenetic analysis or understanding evolution are separate issues. So when interpreting the literature in which character states are optimised on a tree, one should bear in mind the problems surrounding the delimitation of states (e.g. Stevens 2000, 2006b). Furthermore, studies have rather unsurprisingly, perhaps, but importantly shown that dividing the one character into different sets of states may yield differing ideas of evolution of that character (e.g. Lamb Frye & Kron 2003; Hibbett 2004).

As one might expect, with increasing knowledge of development, etc., delimitation of states and characters does not necessarily become easier. Thus Buzgo et al. (2004) and Matthews and Endress (2005) show how hard it can be to delimit states or even to distinguish between e.g. prophylls and other floral structures, Peney et al. (2005) found that not all monosulcate pollen grains in monocots have the same developmental pathway, and therefore such pollen might not have the same ancestral state, while Reeves and Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in Lamiales and Solanales were different and Serna and Martin (2006) described similar problems with the development of hairs in Arabidopsis and Antirrhinum and Solanaceae.

Endress and Matthews (2006a) emphasize the importance of tendencies and developmental constraints when thinking about characters of clades within the rosids - many characters, or character combinations, in these clades occur in a rather sporadic fashion within the clade and apparently notably less frequently outside it. There are a number of other examples in the order pages here, ranging from the features enclosed by parentheses in groups above the level of family, e.g. "(cuticular waxes as aggregated rodlets)" for the commelinids to the discussion of ovular features in the asterids (see Cornales page), although I have kept few of these features in this page. One of the most striking examples is the distribution of N-fixation, restricted as it is to a monophyletic group of four clades, although it has arisen seven or more times independently within that clade (e.g. Clawson et al. 2004). More concrete examples of work that bears on the issue of tendencies include the findings that flowers of polysymmetrical Arabidopsis have genes like TCP1 that are expressed asymmetrically during early development, and TCP1 is a probable orthologue of the well-known CYC gene of Antirrhinum (Cubas et al. 2001; Costa et al. 2005; etc.). Parallelism might build on this underlying morphologically cryptic monosymmetry, even if details of the genetic mechanisms causing the monosymmetry evident in particular groups may be different (Reeves & Olmstead 2003; Cubas 2004 see above). It has also been suggested that for some secondary metabolites in particular there is aquisition of the capability to synthesise a particular metabolite; this is then switched off, but not lost - and is sometimes reacquired again (e.g. Wink 2003; Liscombe et al. 2005). Hence the rather spotty distribution of some of these metabolites when considered in the context of phylogenies. As a non-botanical example - but a rather nice one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been considerable parallelism (40-65 or more independent origins) in the evolution of eyes in metazoans. However, Pax 6 seems to be a master control gene that is involved in eye formation perhaps throughout all bilateralians (e.g. Gehring & Ikeo 1999; Erwin & Davidson 2002). When thinking of the association of camplylotopy and micropyle type, or integument thickness and vascularization of the integument (Endress & Matthews 2006a), one can imagine fairly simple developmental preconditions being operative: as Endress and Matthews (2006a) note, it is difficult to imagine vascular tissue developing in an integument only two cells thick. Changes here would be loosely correlated, if morphologically linked. However, the strongly correlated changes noted by Givnish et al. (2005) are ecologically linked but are presumably morphologically/developmentally independent. When there seem to be characters evolving more or less together, one needs to test to see if the changes are concentrated on certain branches of the tree (Maddison 1990; Maddison & Maddison 2000).

I use Remane's three main criteria of "homology", special properties, position, and intermediates, when determining the basic similarity of structures on different organisms. Even these criteria may not yield an unambiguous answer as to what a structure "is", even given a solid phylogeny. As Endress (2005c) observed, a number of features - position, function, development, shape, anatomy, histology, gene activity, and relationships to other taxa that clearly have petals - can be used to distinguish a petal (for example) from other floral structures, so if a petal does not have one of these features, is it thereby not a petal? There is rampant parallelism in seed plants, yet, as mentioned elsewhere, the terms we use to describe seed plants do not replect this; to reform terms so that they relected phylogeny/synapomorphies would cause an immense amount of grief! Given our current understandings of both phylogenies and characters, evolution of some characters in which we are interested seems to be very labile indeed (e.g. see Soltis et al. 2005b), and I have tried to be very cautious when talking about character evolution.

Finally, far too many features are still described as encompassing alternatives or a range of variation. This is perhaps an unlikely situation for apomorphies for major clades, although less so for - say - ecological preferences at the species level (Hardy 2006).

As an example of the cumulative effect of some of these problems, below I suggest two sets of possible apomorphies for Dipsacales, the first (A) is that if [Columelliaceae + Desfontaineaceae] is their sister group and includes a characterisation of that larger group, the second (B) is the one given in the Dipsacales page, which simply represents the characters common to Dipsacales and is largely innocent of any considerations of their possible outgroups.

A. [Columelliaceae + Desfontaineaceae] + Dipsacales: Group I secoiridoids +; cork deep-seated; nodes ?; pericyclic fibers 0; petiole bundles arcuate; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; flowers moderate in size, G inferior, (many ovules/carpel), style long; K persistent in fruit; exotestal cells enlarged, variously thickened and lignified.

Dipsacales: True tracheids +; nodes 3:3; buds with scales; nectaries as multicellular hairs on corolla, pollen grains trinucleate; testa vascularised, (exotestal cells palisade).

But of course even these characterisations may be strongly affected by the features of the clade sister to any [[Columelliaceae + Desfontaineaceae] Dipsacales] grouping...

B. Dipsacales: Group I secoiridoids +; true tracheids +; nodes 3:3; petiole bundles arcuate; buds with scales; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; nectaries as multicellular hairs on corolla, pollen grains trinucleate; K persistent in fruit; testa vascularised, exotestal cells enlarged (palisade), variously thickened and lignified.

Also included in the synapomorphy hierarchy are brief indications of the sizes and distributions of families and subfamilies, and usually also features by which families can be recognised. If you click on a family name that is highlighted, you will be taken to that family on the relevant ordinal page. You can get back here from any ordinal page by clicking on the highlighted "Apomorphies" immediately after the ordinal name.

SEED PLANTS

Plant woody, evergreen; true roots present; nicotinic acid metabolised to trigonelline; lignins rich in guaiacyl units; ectophloic eustele [in stem] +, endodermis 0; stem xylem endarch, root xylem exarch; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem differentiating internally, phloem externally]; wood homoxylous, tracheids +; sieve tube/cell plastids with starch grains; phloem fibers +; nodes 1:1; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral; plant heterosporous; true pollen [microspores] +, mono[ana]sulcate, pollen exine and intine homogeneous, ovules unitegmic, crassinucellate [megasporangium eusporangiate], megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development endo/exosporic, gametes two, with cell walls; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", embryo straight, endoscopic, short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication, mitochondrial nad1 intron 2 present.

GYMNOSPERMS  Back to Main Tree

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 +; pollen endexine lamellate, pollen chamber + [developing by breakdown of nucellar cells]; pollination droplet +, fertilisation 4-6 months or more after pollination, pollen tube breaks down sporophytic cells, two prothallial cells and two male gametes; female gametophyte 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 nuclear divisions, massive suspensor formed; gametophyte persists in seed; two copies of LEAFY gene.

CYCADALES Dumortier

Roots and stems with contractile tissue; ß-methylamino-L-alanine and compounds producing methylazoxymethanol +, mucilage copious; association with Nostoc or Anabaena in apogeotropic coralloid roots, root hairs 0; primary thickening meristem +; wood manoxylic; 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 in determinate cones, many abaxial microsporangia/sporophyll, dehiscing by the action of the epidermis [exothecium]; megasporophylls with terminal sterile portion; pollen tube usually unbranched, growing away from the ovule, one prothallial cell, spermatogenous cells delimited by circular anticlinal wall, zooidogamy, male gametes with cell walls, released from the swollen proximal part of the tube, flagellae numerous; seeds with sarcoexotesta, scleromesotesta, and ± degenerating endotesta; 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: axillary buds +.

GINKGOALES Gorozh.

VAM present; biflavones, non-hydrolysable tannins +, lignins lacking syringaldehyde [Mäule reaction negative]; wood pycnoxylic; side wall pits in tracheids with tori; compression wood +; leaves with two traces, venation dichotomising, open; plant dioecious; 2 pendulous microsporangia/microsporophyll, dehiscing by the action of the hypodermis [endothecium], exine thin [<2 µm across]; 2(-4) terminal erect ovules/podium, with basal collar, only inner integument vascularised; pollen tube branched, growing away from the ovule, prothallial cells 2, spermatogenous cells delimited by circular anticlinal wall, zooidogamy, male gametes with cell walls, released from the swollen proximal part of the tube, flagellae numerous, pollen tube penetrates between sporophytic cells; 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 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; pollen tube unbranched, growing towards the ovule, prothallial cells 2, gametes non-motile, released from the distal end of the tube, siphonogamy; seed coat 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 0sieve cells with nacreous walls, phloem fibers 0; ; sieve tube plastids with protein fibers; 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; pollen intectate, granulate; pollen germinates on ovuliferous scale [?Araucaria], 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; seed arillate. - 1/ca 5. The Philippines (N. Luzon) to Australia (Tasmania) and New Zealand.

• Phyllocladus has phylloclades, flattened, photosynthetic stems; these bear highly reduced, scale-like leaves, and it is the axils of these leaves that the reproductive structures are found.

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 bladders; prothallial cells 0; sperm cells produced.

SCIADOPITYACEAE Luersson - Leaves reduced to brown scales, short shoots with photosynthetic phylloclades; microsporophyll with flattened apical expansion; 7-9 ovules/ovuliferous scale. - 1/1. C. and S. Japan.

• Sciadopityaceae have leaves borne in loose whorls at the ends of the innovations. Not only are these leaves borne axillary to reduced leaves, as in Pinaceae, but they appear to be highly modified stem structures.

Cupressaceae + Taxaceae: cone scales opposite.

CUPRESSACEAE Bartling - 29/140. Northern hemisphere, more scattered in south temperate region, also NE Africa.

TAXACEAE Berchtold & J. Presl - Plant dioecious; pollen inaperturate; ovule solitary, on shoot in axils of vegetative leaves; male gametes unequal in size. - 6/30. Scattered in the Northern Hemisphere, esp. South East Asia, also New Caledonia.

GNETALES Luersson

Lignins with syringaldehyde [Mäule reaction positive]; stem apex with tunica/corpus construction; roots diarch; vessels + [from circular bordered pits], both fiber tracheids and tracheids +; side wall pits in vessels with tori; mucilage cells +; leaves with two traces; stomata syndetochelic; leaves opposite, joined at the base, with collateral buds; strobilus compound, bracts opposite; plant dioecious, but micro- and sterile megasporangium-bearing structures at least sometimes closely associated; 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"]; male gametophyte with two prothallial cells, a sterile cell, tube nucleus, and sperm cells, pollen tube unbranched, growing towards the ovule, sperm cells binucleate, siphonogamy, gametes non-motile, released from the distal end of the tube, both fuse with female gametes; secondary suspensor developing from upper embryonal tier, no primary suspensor; germination epigeal; 5' end of the inverted repeat extended. - 3 families, 3 genera, 96 species.

EPHEDRACEAE Dumortier - Xeromorphic; nodes 1:2; leaves reduced, or at least without a lamina; microsporangiophores with 2-8 synangia, each with 2(-4) sporangia, dehiscence porose, pollen lacking a colpus, exine shed on germination; archegonia exposed at base of deep pollen chamber; each nucleus of free-nuclear stage forms an embryo; each nucleus of seed with papillae on the inner side of the outer covering; n = 7. - 1/65. North (warm) temperate, W. South America; drier habitats.

• Ephedraceae are rather small, shrubby, much-branched, xeromorphic plants that may be readily recognised by their green, photosynthetic stems, often reduced opposite leaves and small cones borne along the shoots.

Gnetaceae + Welwitschiaceae: torus:margo pits 0; branched sclereids +; stomata syndetocheilic; 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.

• Gnetaceae are woody, evergreen plants with opposite, angiosperm-like leaves whose bases surround the stem; axillary to the leaves are a series of strobili.

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.

• Welwitschiaceae can immediately be recognised by the two, long leaves that ensheath the stem; these become curled and are dead and torn at the apices. The plant has no trunk as such, but the stem becomes quite massive.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway, lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; cork cambium in root deep-seated, trichoblasts [differentiated hair-forming cells] 0, origin of epidermis with no clear pattern [probably from inner layer of cap]; stem with 2-layered tunica-corpus construction; cork cambium in stem superficial; wood fibers and wood parenchyma +; circular bordered pits lacking margo and torus; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with sieve plate, companion cells from same mother cell that gave rise to the sieve tube and P proteins, plastids with starch grains alone; nodes unilacunar; stomata paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, A many, development centripetal, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, thecae dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther, tapetum glandular, binucleate, microsporogenesis successive, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous, endexine compact, lamellate only in the apertural regions, pollen tube elongated, siphonogamy, gametes 2, with cell walls, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, ovules marginal, anatropous, bitegmic, micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin, chalazal, female gametophyte ?type, stylodium short, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm ?diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio; germination hypogeal, sympodial seedlings/young plants; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA/PHYC gene pairs.

Possible angiosperm synapomorphies are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, partly because many taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied and the immediate outgroup to angiosperms is also uncertain.

AMBORELLALES Melikian, A. V. Bobrov & Zaytzeva  

Nodes 1:1; plant dioecious; hypanthium +, A sessile, vascular bundle branched towards apex, microsporogenesis successive, pollen ektexine granulate, 1 ovule/carpel, embryo sac monosporic, spore chalazal, 8-nucleate, bipolar; fruit of drupelets; endosperm triploid. - 1 family, 1 genus, 1 species.

AMBORELLACEAE Pichon - 1/1. New Caledonia.

• Amborellaceae are sprawling shrubs with apparently two-ranked, exstipulate leaves; the margins are both serrate and rather undulate. The plant is dioecious and the flowers are small, with an undifferentiated perianth. The staminate flowers have sessile anthers and the carpellate flowers a few carpels that develop into drupelets with pock-marked stones and pockets of almost resinous substances.

NYMPHAEALES + AUSTROBAILEYALES + [MONOCOTS + CERATOPHYLLALES] + [CHLORANTHALES + MAGNOLIIDS + EUDICOTS]: vessels +, elements with scalariform perforation plates; pollen tectate-columellate, tectum reticulate [perforated]; nucleus of egg cell sister to one of the polar nuclei; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

NYMPHAEALES Dumortier  

Aquatic herbs; hydrolyzable [ellagi]tannins +; 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 copious [starchy], suspensor 0, embryo broad; seedling/young plant monopodial; intergenic inversion in chloroplast inverted repeat. - 3 families, 10 genera, 74 species.

Cabombaceae + Nymphaeaceae: leaves involute, peltate, 2ndary veins palmate; 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; exotesta palisade, anticlinal walls sinuous.

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

• Cabombaceae are rather small-flowered waterlilies with few ovules or seeds in each carpel; they have floating stems and all the flower parts are free.

NYMPHAEACEAE Salisbury - 6/58. World-wide.

Nupharoideae Ito - K 5-14, pollen spiny. - 1/ca 10. North Temperate.

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. - 5/48. World-wide.

• Nymphaeaceae are the quintessential waterlilies, with large flowers, usually many free perianth parts and stamens, and many ovules in each carpel, the placentation usually being clearly laminar. There is no stem, the plants essentially being rooted, aquatic, rosette plants.

AUSTROBAILEYALES + [MONOCOTS + CERATOPHYLLALES] + [CHLORANTHALES + MAGNOLIIDS + EUDICOTS]: ethereal oils in spherical idioblasts [lamina and P ± pellucid-punctate]; tension wood 0; 12BP [4 amino acids] deletion in P1 gene.

AUSTROBAILEYALES Reveal  

Tiglic acid +; nodes 1:2; P not obviously in 3's, extragynoecial mucilaginous compitum +, outer integument 5-7 cells thick; fruit a berrylet; mesotestal cells ± sclerotic. - 3 families, 5 genera, 100 species.

AUSTROBAILEYACEAE (Croizat) Croizat - Flowers with internal staminodes. - 1/2. Australia.

• Austrobaileyaceae are lianes with opposite and entire leaves and large, pendulous flowers. These have spreading petals, many stamens with broad filaments, and internal staminodes. The fruit consists of more or less ellipsoid berrylets.

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 (J. R. Perkins & Gilg) Gibbs - Rays 6-9-seriate; flowers small, carpels 1-2, 1 ovule/carpel, micropyle bistomal. - 1-2/6. New Guinea and S.E. Australia to Fiji.

• Trimeniaceae are trees or lianes with opposite, serrate to entire leaves that often have pellucid glands and widely spreading and quite close secondary veins. The wind-pollinated flowers are small, there is usually only a single carpel with a single ovule, and the fruit is a berrylet.

CERATOPHYLLALES + CHLORANTHALES + MAGNOLIIDS [MONOCOTS + EUDICOTS]: carpels plicate; embryo sac bipolar, 8 nucleate; endosperm triploid.

CERATOPHYLLALES Bischoff  

Aquatic herb; ethereal oils 0; roots 0; vessels 0; nodes?; stomata 0; leaves opposite, margins spiny-toothed; plant monoecious, flowers extraxillary, alternating with leaves; P not vascularised, A extrorse, connective apically produced, tapetum amoeboid, pollen 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] 0, suspensor 0, embryo large. - 1 family, 1 genus, 1-2+ species.

CERATOPHYLLACEAE Gray - 1/2 or more.  World-wide.

• Ceratophyllaceae may be recognised by their whorled, spiny-toothed, often dichotomously-branched leaves; their small, sessile flowers (even the perianth and connective have spiny teeth); and their distinctive, spiny, achenial fruits.

CHLORANTHALES + MAGNOLIIDS [MONOCOTS + EUDICOTS]: benzylisoquinoline alkaloids +.

CHLORANTHALES J. F. Leroy  

?Neolignans, sesquiterpenes +; rays 6-10-seriate; nodes often swollen, ?type; leaves opposite, toothed, teeth with lateral vein and others; stipules +; flowers very small, monosymmetric, parts whorled; P 3 or 0, A 1, G 1, ± inferior, ascidiate, postgenital fusion by secretion, 1 apical atropous ovule/carpel; fruit fleshy; endotesta palisade, lignified, crystalliferous. - 1 family, 4 genera, 75 species.

CHLORANTHACEAE Sims - 4/75. Tropics and subtropics, not Africa (Madagascar - Chloranthus only).

• Chloranthaceae are aromatic plants that may be recognised by their opposite, serrate leaves with more or less sheathing stipules and often strongly swollen nodes that may collapse on drying. The flowers are very small and are often borne on branched inflorescences. The wood is soft.

MAGNOLIIDS [MONOCOTS + EUDICOTS]: P more or less whorled, 3-merous [possible position], A whorled, carpel fusion by congenital occlusion.

MAGNOLIIDS: (neolignans +); leaf margins entire; A many, spiral [possible position here], extrorse, antipodal cells ephemeral, hypostase +, nucellar cap +, raphal bundle branches at the chalaza.

MAGNOLIALES + LAURALES: cuticle waxes as annularly-ridged rodlets, palmitol the main wax.

MAGNOLIALES Bromhead  

Sesquiterpenes +; 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; P 3, connate; A whorled, connate, 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.

• Myristicaceae are easily recognised: the plant has red sap, the terminal bud is long, the leaves are usually 2-ranked, the petioles are often short and the blades are often glaucous below. The blades are often much damaged by insects, they are brittle when dry, and the venation is inconspicuous. Growth is fairly stereotyped: the main axis is orthotropic and bears pseudoverticillate, radially-departing branches at the end of each flush. The plants are dioecious and the rather small flowers have a connate, 3-merous perianth, connate stamens, and a single carpel. The single-seeded, dehiscent fruits are often quite large and the seed is ruminate and arillate.

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 +; stipules surrounding stem; flowers terminal; receptacle elongated, micropyle bistomal. - 2/227. The Americas (but not W. North America), and South East Asia to Malesia.

• Magnoliaceae may be recognised by their entire or lobed leaf blades that are often more or less glaucous below and have large stipules that entirely surround the stem but are open on the side opposite the petiole. The flowers are rather large, usually single, and have an elongated receptacle bearing many petals, stamens and carpels that are obviously spirally arranged.

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.

• Degeneriaceae may be recognised by their spiral, entire, estipulate leaves and large, axillary flowers with many tepals and a single 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.

• The 2-ranked, estipulate leaves, the peltate scales that cover the plant, the flowers with many petals, and the more or less syncarpous fruits with laterally compressed seeds allow one to recognise the family. The crushed leaves are very aromatic.

Eupomatiaceae + Annonaceae: sieve tube plastids with protein crystalloids and starch; rays 8-15-seriate; petiole bundles arcuate; 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.

• Eupomatiaceae may be recognised by their 2-ranked, exstipulate, aromatic leaves and their fairly large, calyptrate flowers with stamens sourrounding many flat, gland-bearing staminodes that arch over the gynoecium. The receptacle is concave and bears many carpels tightly packed together with sessile but basically decurrent stigmas. The stamens and staminodes fall off the flower together, littering the ground beneath the plant with what look rather like sea anemones.

ANNONACEAE Jussieu - Flowers with open development; A whorled, filaments with a single vein; endotestal plug +; tegmen crushed, ruminations irregular. - 129/2220. Largely tropical.

Anaxagorea - Uniseriate hairs terminate in a rounded cell; fruit a follicle; endotesta aerenchymatous, tegmen alone involved in ruminations, with idioblastic oil globules. - 1/21. Tropical America, also Sri Lanka to West Malesia.

The rest - Stamens with prominent ± truncate connective, internal staminodes 0. - 128/2200. Predominantly lowland tropics.

• Annonaceae are easily recognised, even when sterile. The twigs often dry blackish, the bark is conspicuously fibrous (when cut tangentially, there is a reticulum), the rays are broad, the young stems are often zig-zag, and the ± aromatic leaves are entire and 2-ranked. The obliquely-pointing bud has clearly conduplicate leaves. The flowers are nearly always pendulous. Their development is more or less open, and they have fleshy tepals usually in three whorls of three and numerous short stamens with a tough, truncate connective. The seeds are quite large and most have a regularly ruminate endosperm.

LAURALES Perleb

Sieve tube plastids with protein crystalloids; nodes 1:?; petiole bundle(s) arcuate; leaves opposite; inflorescence ± cymose; hypanthium +, inner staminodia +, ovules basal, archesporium multicellular, stylodium long; fruitlets 1-seeded, indehiscent, hypanthium persistent; mesotesta crushed, endotesta tracheidal [seed endotestal], tegmen crushed; embryo long; duplication of the PI gene. - 7 families, 91 genera, 2858 species.

CALYCANTHACEAE Lindley - Stem with four inverted cortical bundles; P many, spiral, no double fertilisation [endosperm develops autonomously]. - 5/11. China, North America, N.E. Australia.

1. Idiospermum - G 1-2(-3), outer integument 12-15 cells thick, micropyle bistomal, nucellar beak +; cotyledons (3-)4, massive, peltate. - 1/1. N.E. Australia.

2. The rest - Pollen euatorially and vertically disulcate, G 5-35, outer integument 5-6(-8) cells thick; cotyledons spirally twisted. - 4/10. China, North America.

• Calycanthaceae can be recognised by their opposite, entire, exstipulate leaves that smell aromatic. The cortical bundles can be seen with a lens when the stem is cut, and they may also be visible externally as four, raised lines along the stem. The terminal bud often aborts. The large flowers have a bract-covered hypanthium; the inner members of the perianth secrete nectar or bear food bodies. The achenial fruits are attached to the inside of the persistent hypanthium.

Siparunaceae + Atherospermataceae + Gomortegaceae + Monimiaceae + Hernandiaceae + Lauraceae: vessel elements with scalariform perforations; hippocrepiform sclereids in pericycle; leaves with rather distant teeth, one vein entering opaque, persistent glandular cap; flowers rather small, A whorled, stamens with paired nectaries/glands at base, anthers bisporangiate/unithecal, valvate, tapetum ?, pollen inaperturate, ± spinulose, 1 ovule/carpel.

Siparunaceae + Gomortegaceae + Atherospermataceae: acicular crystals +; hypanthium closed by roof, infratectum columellar; embryo very small.

SIPARUNACEAE (A.-L. de Candolle) Schodde - A bisporangiate/unithecal, with a single slit, integument 0. - 2/75. Tropical America, W. Africa.

• Siparunaceae may be recognised by their opposite, serrate, estipulate leaves with rather prominent brochidodromous venation; the wood lacks broad rays and the hairs are often stellate or fasciculate. The anthers lack basal glands.

Gomortegaceae + Atherospermataceae: bud scales +; sieve tube plastids also with fibrils; outer, not inner, A staminodial.

GOMORTEGACEAE Reiche - G inferior, ovule apical, hemianatropous, stigmatic branches erect; embryo large. - 1/1. C. Chile.

• Gomortegaceae are aromatic plants with opposite, estipulate, entire leaves. The flowers are small, with staminodes outside the stamens, the ovary is inferior and syncarpous, and the single-seeded fruit is drupaceous.

ATHEROSPERMATACEAE R. Brown - Stomata anomocytic; fruit achenial, plumose, hypanthium woody. - 6-7/16. New Guinea to New Zealand and New Caledonia, Chile.

• Atherospermataceae have opposite, coarsely serrate leaves with brochididromous venation. The flowers are quite small, the fruit is dry, and the fruitlets plumose and achenial.

Monimiaceae + Hernandiaceae + Lauraceae: A whorled, columellae foot layer and endexine 0, ovule apical.

MONIMIACEAE Jussieu - 22/200. Tropical, but esp. Australasia.

• Monimiaceae are recognisable by their opposite, rather distantly serrate leaves which have prominent brochidodromous venation on the lower surface, and their twigs, which have broad rays and are often flattened below the somewhat swollen nodes; some taxa have "laticiferous threads" (Keller 1996). In fruit, the well-developed fleshy hypanthium and/or receptacle is conspicuous; this may enclose the drupelets and it then splits irregularly.

Hernandiaceae + Lauraceae: primary stem ± with vascular cylinder; vessel elements with simple perforations; mucilage cells +; sieve tube plastids also with starch; leaves spiral (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 - 3/44. Tropical, esp. Madagascar and Indo-Malesia.

Gyrocarpoideae Pax - 2/10. Pantropical, esp. America.

LAURACEAE Jussieu - Leaves entire; fruit a drupe. - Ca. 50/2500. Pantropical (temperate), lowland to montane.

• The family may be recognised by its aromatic smell, nearly always entire spiral or opposite (very rarely indeed 2-ranked) leaves the blades of which are often quite thick (they are coriaceous when dry) and with a glaucous undersurface and rather steeply ascending secondary veins. The twigs are often more or less ridged, and branching is often on the current flush - even in temperate taxa there is often a mixture of proleptic and sylleptic branching on the same plant. The flowers are small, trimerous, and have valvate anthers; the fruit has a single, large seed that lacks endosperm and the pedicel - and sometimes also tepals and hypanthium - is often ± swollen and conspicuous. Some Lauraceae have a growth pattern similar to that of Myristicaceae, with whorled branches and periodic growth, but the leaves on the branches of Lauraceae are usually spiral, rarely opposite, while in Myristicaceae (and Annonaceae) the leaves are distichous.

CANELLALES + PIPERALES: sesquiterpenes, 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.

• Canellaceae are aromatic evergreen shrubs or trees that may be recognised by their rather thick, exstipulate leaves with entire margins and flowers with three outer sepalline tepals, connate stamens with extorse anthers, and ovary with parietal placentation. The fruit is a berry.

WINTERACEAE Lindley - Rays 10+-seriate; K calyptrate. - 4-7/60-90. Montane tropics, not mainland Africa.

Taktajanioideae Leroy - Stomata anomocytic; G [2], placentation parietal. - 1/1. Madagascar.

Winteroideae - 3-6/60-90. New Guinea to New Zealand and New Caledonia, few Borneo and the Philippines and South America.

• Winteraceae are often rather small evergreen trees or shrubs that can be recognised by their aromatic and/or peppery taste and their spiral, entire leaves that are often glaucous below. Their flowers have a more or less obviously calyptrate calyx although the other parts are nearly always free. The fruit is often a berrylet.

PIPERALES Dumortier

Herbs; vessel elements with simple perforations; starch grains compound; primary stem with distinct bundles; vessel elements with simple perforations; wood with broad rays; nodes often swollen; stomata ?; leaves 2-ranked, 2ndary veins palmate; flower parts whorled, P and A in 3's, G occlusion?; seed ± tegmic. - 5 families, 17 genera, 2509 species.

Hydnoraceae + Lactoridaceae + Aristolochiaceae: embryo undifferentiated.

HYDNORACEAE C. Agardh - Echlorophyllous root parasites; leaves 0; flowers arising endogenously from roots, large, P uniseriate, connate, valvate, thick and fleshy, A = adnate to and opposite P, A sessile, connate, polythecate, microsporogenesis successive, 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.

• Hydnoraceae are leafless and echlorophyllous parasites that can be recognised by their large flowers with a single, thick, basally connate, valvate perianth whorl that is brownish and cracked outise and coloured inside, stamens opposite the perianth lobes, and inferior ovary; the fruit contains many small seeds.

Lactoridaceae + Aristolochiaceae: stomata anomocytic; prophyll adaxial; inflorescence cymose, nucellar cap +, micropyle endostomal.

LACTORIDACEAE Engler - Stipules intrapetiolar; pollen in tetrads, saccate. - 1/1. Chile, the Juan Fernandez Islands.

• Lactoris can be recognised by its small, 2-ranked leaves with intrapetiolar stipules; the prophylls are adaxial. The flowers are small and have three imbricate perianth members and three free carpels.

ARISTOLOCHIACEAE Jussieu - P connate, nectaries or secretory hairs on tube, A ± connate, pollen inaperturate, ektexine semitectate-reticulate, ± granular, G ± inferior; endotesta palisade, usu. crystalliferous, exotegmen and layer underneath crossing fibers, endotegmen with reticulate thickenings, tanniniferous; embryo undifferentiated. - Ca 4/480. World-wide, not Arctic.

Asaroideae O. C. Schmidt - Growth sympodial; sieve tube plastids with cuneate protein crystalloids and a large polygonal crystal; inflorescences/flowers terminal; hypanthium +, P = K + C, pollen sulcate. - 2/75. N. Temperate, esp. East Asia.

Aristolochioideae - Viny; growth monopodial; benzylisoquinoline alkaloids +; sieve tube plastids with polygonal protein crystalloids plus starch grains or protein fibers; groups of silicified cells +; serially arranged axillary buds; hairs hooked; inflorescences axillary; floral primordia monosymmetric. - 2/405. Tropics (temperate), relatively less diverse in Africa (inc. Madagascar), few in N. Australia.

• Aristolochiaceae are more or less herbaceous plants (they are quite often vines) that may be recognised by their adaxial prophylls, exstipulate leaves with palmate venation and entire margins, and 3-merous flowers with extrorse stamens. The ovary is usually inferior and the perianth uniseriate and connate. Only Aristolochia has monosymmetric flowers, only Saruma a perianth differentiated into sepals and petals and a largely superior gynoecium of free carpels.

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. - 5-8/2015. Pantropical.

Piperoideae - Vascular bundles in more than one ring. - 4/2000. Pantropical.

Peperomioideae - Vascular bundles scattered; pollen inaperturate, G 1, ovule unitegmic [2 cell layers across], tenuinucellate, embryo sac 16-nucleate, chalazal nuclei fuse, stigma penicillate; seed endotestal, endosperm cellular. - 1-4/1000. Pantropical, esp. America.

• Piperaceae may be recognised vegetatively by their herbaceous habit and/or their separate vascular bundles in the stem, their swollen nodes, and their soft and/or fleshy leaves that are often cordate at the basde. The minute flowers lack a perianth and are often borne in dense spikes. The individual flowers are subtended by peltate to clavate bracts. Some species of Piper look remarkably like Araceae!

SAURURACEAE Martynov - A introrse. - 4/6. North Temperate.

• Saururaceae may be recognised by their often aromatic and rather fleshy leaves with broad bases and palmate venation and by their spicate inflorescences that usually have a large, basal inflorescence bract. The flowers are small, without a perianth, but there are clearly three to four carpels.

MONOCOTS + EUDICOTS: (stamens opposite [two whorls of] P).

MONOCOTYLEDONS: Herbaceous, plant sympodial; non-hydrolyzable tannins [(ent-)epicatechin-4] +, ellagitannins, neolignans 0, hemicelluloses as xylans; root apical meristem?; root epidermis developed from outer layer of cortex; trichoblasts in vertical files with proximal cell smaller or hypodermal cells dimorphic; cork cambium in root [uncommon] superficial; root vascular tissue oligo- to polyarch, medullated, lateral roots arise opposite phloem poles; tunica 2-layered [?sampling]; primary thickening meristem +; vascular bundles in stem scattered, (amphivasal), closed [no interfascicular cambium developing]; vessels in root with scalariform and/or simple perforations; vessels in stems and leaves 0; sieve tube plastids with cuneate protein crystals alone; epidermis with bulliform cells [?this level], stomata paracytic [cell divisions oblique]; leaves not differentiated into petiole plus lamina, main venation parallel, developing both acropetally and basipetally from the base and converging towards the apex, intermediate [and other] veins basipetal from apex, endings not free, (margins with spiny teeth), Vorläuferspitze +, base sheathing, sheath open, colleters [intravaginal squamules] +; inflorescence racemose; flowers 3-merous, polysymmetric, pentacyclic, T (each with three traces), in two whorls, median member of outer whorl abaxial, members of whorls alternating, similar, [pseudomonocyclic, each providing a sector for the T tube when present], A = and opposite each T member (primordia associated), anther and filament sharply distinguished, microsporogenesis successive, G [3], development?, opposite outer tepals [thus median member abaxial], placentation axile, style hollow, short; fruit a loculicidal capsule; seed testal; embryo long, cylindrical, cotyledon 1, terminal, plumule lateral; seedling phanerocotylar, primary root unbranched, adventitious roots numerous, hypocotyl short, (collar rhizoids +), cotyledon with a closed sheath, unifacial, assimilating and haustorial; duplication producing monocot LOFSEP gene.

Some features that are likely to be synapomorphies - almost whatever the immediate sister taxon to monocots might be - are in bold. However, if Ceratophyllaceae are sister to monocots, synapomorphies like the herbaceous habit, absence of vascular cambium, etc., will move down a node.

ACORALES Reveal  

Inflorescence a spadix, ebracteate, with spathe; ovules atropous; endosperm copious, perisperm +. - 1 family, 1 genus, 2-4 species.

ACORACEAE Martinov - Leaves equitant, isobifacial; peduncle with two separate vascular systems; anther thecae with confluent dehiscence, pollen sulcus lacking ectexine. - 1/2-4. E. America to East and South East Asia.

• Acoraceae are recognisable by their sweetly-smelling, two-ranked, isobifacial leaves and their densely spicate inflorescence overtopped by and appearing lateral to the leaf-like spathe. The flowers are small, perfect and pentacyclic.

ALISMATALES + PETROSAVIALES + [DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: ethereal oils 0; leaf ptyxis variants of supervolute-curved; endothecium develops directly from undivided outer secondary parietal cells, endexine 0, carpels plicate, (septal [epithelial] nectaries +); endosperm nuclear/helobial.

ALISMATALES Dumortier  

Starch grains pteridophyte-type, amylophilic; A extrorse, tapetum amoeboid, cells uninucleate, carpels with completely unfused canals, styles separate; endosperm helobial; embryo large; seedling with hypocotyl and root well developed. - 14 families, 166 genera, 4490 species.

ARACEAE Jussieu - Inflorescence terminal, densely spicate [spadix], subtended by an inflorescence bract [spathe], bracts 0; pollen endexine spongy. - 106/4025. Mostly tropical.

Gymnostachydoideae + Orontioideae: ovules atropous.

1. Gymnostachydoideae Bogner & Nicolson - Nucellus not fully covered by integument. - 1/1: Gymnostachys anceps. E. Australia.

2. Orontioideae Mayo, Bogner & Boyce - 3/6. Temperate East Asia, W. and E. North America.

3. Lemnoideae Engler - Thallus-like, stemless, floating aquatic herbs; collenchyma and bundle fibers 0; vessels 0; P 0, A 1, monothecal, or 2, pollen ulcerate, spiny, G 1, embryo sac Allium-type, stigma funneliform; fruit an achene [sort of]; seed operculate; endosperm cellular, starchy, embryo undifferentiated. - 5/37. World-wide.

4. Pothoideae Engler

4a. Potheae Engler - 4/900. Tropical America, Madagascar to South and Southeast Asia, Malesia and N.E. Australia.

4b. Monstereae Schott - 12/360. Tropical South and Southeast Asia to the Pacific, South America (Africa).

5. Lasioideae Engler - Inflorescence flowering basipetally; pollen sulcus with ectexine lamella and thick bilayered endexine [outer: flakes or lamellae; inner: spongy]. - 10/58. India and Southeast Asia to the Pacific, also Africa (Lasimorpha).

Calloideae + Aroideae: ?

6. Calloideae Endlicher - P 0, pollen dicolpate. - 1/1: Calla palustris. Circumboreal.

7. Aroideae - Plants monoecious (dioecious); spathe differentiated into tube plus blade, pollen extruded in strands. - 70/2670. Tropical and warm temperate.

7a. Zamioculcadeae Engler - Leaves compound, petiole geniculate; staminate flowers: pollen zonasulcate, columellae winding, forming a sort of internal tectum as well as the external tectum, endexine lamellate, intine thin. - 2/6. Africa, Kenya to Natal.

7b. The rest - Laticifers +; P 0, A connate, connectives thick, pollen inaperturate, ektexine thin, often lacking sporopollenin, endexine thick, spongy, intine massive. - 68/2665. Tropical and warm temperate.

• Araceae are herbs with leaves that are usually divided into a petiole and expanded blade. They have a distinctive inflorescence of a more or less petaloid spathe plus a spadix made up of densely-packed, sessile, ebracteate flowers - in many taxa the spadix has a large, terminal, sterile part. The fruit is a berry.

Tofieldiaceae + [Hydrocharitaceae + Scheuchzeriaceae groups]: ?

TOFIELDIACEAE Takhtajan - Leaves equitant, isobifacial; flowers subtended by a "calyculus"; A introrse to latrorse, tapetum secretory, microsporogenesis simultaneous [tetrads tetrahedral], pollen disulcate, nectaries +, ovules unitegmic. - 3-5/27. S.E. U.S.A., N.W. South America, N. temperate.

• Tofieldiaceae can be recognised by their equitant, isobifacial leaves; racemose inflorescences in which there is usually a 1-3-membered "calyculus" below the flower; and undistinguished monocot flowers with the tepals usually free and the tips of the carpels each with a clearly separate stylodium. The fruit is usually a septicidal capsule.

Hydrocharitaceae + Scheuchzeriaceae groups: plant aquatic (with floating stems); stem with lacunae; raphides and druses 0; hairs 0; bulliform cells 0 [?this level]; pollen grains trinucleate; endosperm 0; seedling collar and collar rhizoids +.

Hydrocharitaceae + Butomaceae + Alismataceae + Limnocharitaceae : C-glycosyl flavones +; inflorescence scapose, determinate, bracteate; P = K + C, (stamen pairs +), carpels plicate; seeds exotestal.

Hydrocharitaceae + Butomaceae: ?

HYDROCHARITACEAE Jussieu - G inferior; 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.

• Hydrocharitaceae are aquatic plants with petiolate or (usually) undifferentiated leaves. The inflorescence often has two fused (sometimes free) bracts at the base and the ovary is inferior, often with laminar or more or less strongly intruded parietal placentation, and the stigma lobes are bifid.

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

• Butomaceae are aquatic monocots that may be recognised by their long, three-angled leaves and axillary, scapose, umbellate, bracteate inflorescence. The flowers are perfect and there are clearly two distinct perianth whorls although both are petaloid; the fruit is a follicle.

Alismataceae + Limnocharitaceae: plant with latex; stomata with parallel divisions; leaves involute, with pseudopetiole, midrib, cross veins and an apical subepidermal pore; C thin, more or less crumpled in bud, evanescent, A centrifugal, pollen pantoporate, spinose, G free or connate basally, ovules tenuinucellate, embryo sac tetrasporic, nucellar cap +; fruit a follicle; embryo strongly curved, white; x = 7, 8.

ALISMATACEAE Ventenat - 12/81. Pantropical, also temperate.

• Alismataceae are laticiferous, aquatic herbs. Their floating or aerial leaves are petiolate and have a prominent midrib, parallel veins and also cross veins. Their petals are crumpled in bud and their staminate flowers usually have many extrorse anthers and their carpellate flowers have many free carpels. It is obvious from looking at the seeds that the embryo is strongly bent.

LIMNOCHARITACEAE Cronquist - 3/7. Pantropical.

• Limnocharitaceae are laticiferous, aquatic herbs. Their floating or aerial leaves are petiolate and have a prominent midrib and parallel veins as well as cross veins. Their flowers have sepals, small to large but evanescent petals, many stamens (and also staminodes), and carpels with many ovules and laminar placentation. It is obvious that the seeds have a strongly curved embryo.

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.

• Scheuchzeriaceae are small herbs of marshy places with distichous leaves; the open sheaths have auricles at the top and there is a little pore at the tip of the blade. The racemose inflorescence has large, leafy bracts and all the parts of the flowers (except the bases of the carpels) are free from one another.

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

APONOGETONACEAE J. Agardh - 1/43. Old World, esp. South Africa, largely tropical and warm temperate.

• Aponogetonaceae are cormose water plants with petiolate leaves; the sometimes fenestrate blades have a midrib, parallel veins, and strong cross veins; the finest veins are reticulate. The spicate inflorescence has a long scape and the rather small flowers nevertheless have more or less conspicuous tepals.

Juncaginaceae + Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: sulphated phenolic acids +, leucanthocyanins, flavones 0; leaf ± linear, base auriculate, ligulate; P 0 or reduced ["abaxial outgrowth" of A], anthers sessile, pollen inaperturate, nectary 0, carpels with complete postgenital fusion [sampling!].

JUNCAGINACEAE Richard - 4/15. Cosmopolitan.

• Juncaginaceae are herbs of marshy places with more or less unifacial leaves and a scapose racemose or spicate inflorescence; all parts of the flower appear free.

Posidoniaceae + Ruppiaceae + Cymodoceaceae + Zosteraceae + Potamogetonaceae: aquatics; rhizome with endodermis; epidermis chlorophyllous; water pollination; carpels ascidiate [sampling!], 1 apical atropous ovule/carpel; 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. - 1/9. Mediterranean, temperate Australia.

• Posidoniaceae are marine "grasses" that are readily recognisable by the fibrous strands persisting along the monopodial rhizome and the branched-racemose inflorescences.

Ruppiaceae + Cymodoceaceae: leaves serrulate; A 2.

RUPPIACEAE Horaninow - 1/1-10. More or less world-wide.

• Ruppiaceae are submerged aquatic plants that may be recognised by their 2-ranked, serrulate leaves with an obvious midrib and a sheathing base; the internodes are well developed.

CYMODOCEACEAE N. Taylor - 5/16. More or less tropical (to warm temperate), Australia in particular.

Zosteraceae + Potamogetonaceae: leaf with apical pore, (sheath closed); plant mono- or dioecious.

ZOSTERACEAE Dumortier - Anther thecae separate, deciduous, joined by divided connective, pollen filiform. - 2/14. Temperate to subtropical.

• Zosteraceae are sea grasses with leaf-opposed branches and flattened inflorescence axes (spadices) with flowers on the adaxial surface; the inflorescences are enclosed by a spathe. Most taxa have "retinacules", perhaps tepals or bracts, opposite the stamens.

POTAMOGETONACEAE Reichenbach - 7/102. Worldwide, esp. temperate.

• Potamogetonaceae are aquatic plants most of which have more or less petiolate leaves with midrib and cross veins; there is usually a conspicuous basal ligule. The inflorescence is densely spicate and the flowers have small tepals appearing to be borne on the backs of the stamens. The carpels are stipitate, as is particularly clear in Zannichellia are particularly obvious, although that genus does not have a spicate inflorescence.

PETROSAVIALES + [DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: Starch grains simple, amylophobic; druses 0, raphides +; stomata anomocytic; colleters 0; endosperm nuclear.

PETROSAVIALES Takhtajan

Stem with a ring of bundles; sieve tube plastids also with polygonal protein crystalloids; microsporogenesis simultaneous, septal nectaries +; fruit a follicle.

PETROSAVIACEAE Hutchinson* - Bracteoles 0; integumentary obturator +. - 2/4. China, Japan, W. Malesia. 1 family, 2 genera, 4 species.

[DIOSCOREALES + PANDANALES] + LILIALES + ASPARAGALES + COMMELINIDS: Inflorescence type?

[DIOSCOREALES + PANDANALES]: ?

DIOSCOREALES J. D. Hooker  

Vascular bundles in rings; 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 - Leaves equitant, isobifacial. - 4-5/41. N. temperate, Venezuela and Guiana.

• Most Nartheciaceae can be recognised by their equitant, isobifacial leaves and racemose, bracteate inflorescences with small, rather undistinguished monocot flowers that have partly free tepals that persist under the capsular fruit.

Taccaceae + Thismiaceae + Burmanniaceae + Dioscoreaceae: stem with endodermis; leaves spiral; inflorescence cymose; 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.

• Taccaceae are herbs with long-petiolate leaves and a scapose inflorescence with medium-sized, purple flowers that have inferior ovary; long, filiform bracts are mixed in with the flowers and the ovary placentation is parietal.

THISMIACEAE Agardh - Plants ± mycotrophic; stomata 0; leaves reduced to scales; T whorls often differentiated, both well developed, A connate postgenitally into a ring, pollen porate, style connate early in ontogeny; fruit irregularly dehiscent, P usually circumscissilely dehiscent. - 4/31. Widely scattered, (sub)tropical.

• Thismiaceae are small, echlorophyllous herbs that may be recognised by their distinctive flowers. The outer tepals are often rather different from the inner, there is usually an annulus surrounding the broad throat, the stamens are reflexed, and the ovary is inferior. The flowers can look like a little jelly fish, or a group of small fungi, or...

Burmanniaceae + Dioscoreaceae: ?

BURMANNIACEAE Blume - Raphides 0; outer T larger, enclosing the inner, A 3, opposite inner T, thecae separated by connective, tranversely dehiscent, stigma infundibular. - 9/95. Largely tropical, esp. American.

• Burmanniaceae are rather small, often echlorophyllous herbs that may be recognised by their radially symmetrical, often more or less blue flowers with an inferior ovary that is often winged. The outer tepals are rather larger than the inner and there are only three stamens, borne opposite the inner tepals.

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. - 4/870. Largely tropical.

• Dioscoreaceae are more or less herbaceous plants with often rather small and not very brightly coloured flowers; the ovary is inferior and often ridged or winged in fruit. Many are climbers recognised by their spiral, petiolate leaves with pulvini at both ends of the petiole; the leaf base does not surround the stem. The lamina has several strong longitudinal veins and transverse secondary veins, or reticulate fine venation. The flowers have an inferior ovary and are mostly rather small.

PANDANALES Lindley   

Nucellar cap +; embryo minute, endosperm type? - 5 families, 36 genera, 1345 species.

VELLOZIACEAE Hooker - Xeromorphic; 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).

• Velloziaceae are xeromorphic and sometimes tree-like monocots (but the "trunk" is made up largely of adventitious roots) that may be recognised by their 3-ranked leaves with persistent bases and spiny margins. The inflorescences are terminal, although often appearing axillary, and often have one flower. The flowers are rather large with petaloid tepals and a long style; there may be a corona, and the stamens can be many.

Triuridaceae + Stemonaceae + [Pandanaceae + Cyclanthaceae]: septal nectaries 0.

TRIURIDACEAE Gardner - Echlorophyllous mycotrophic herbs; vessels 0; stem with a ring of bundles; stomata 0; inflorescence a raceme; T valvate, carpels free, 1 (2) basal tenuinucellate ovule/carpel. - 8/48. Pantropical.

• Triuridaceae are mostly rather small echlorophyllous mycoheterotrophic saprophytes with small flowers that have pointed tepals and many separate carpels; the latter often have a decidedly irregular surface in fruit.

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

STEMONACEAE Caruel - Leaves with petiole and well-developed fine cross veins; anther connective expanded; seeds with longitudinal ridges. - 4/27. China and Japan to Australia, S.E. U. S. A.

Stemonoideae - Tuber or rhizome with scale leaves; hairs 0; testa several-layered, ridges many cells high. - 3/25. China and Japan to Australia, S.E. U. S. A. (Croomia ).

Pentastemona - Hairs uniseriate; leaves with compound midrib; flowers 5-merous, A 5, connate into a fleshy ring and basally adnate to T and connectives apically adnate to stigmas, pollen inaperturate, atectate; endotesta with massive U-shaped thickenings. - 1/2. Central Sumatra.

• Stemonaceae can be recognised by their elegant leaves with a petiole, sometimes a distinct midrib, and very close to distant longitudinal veins; there are conspicuous cross veins. When dry, the leaves are often very thin. The inflorescences are axillary and the flowers are 4- or 5-merous.

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, with adventitious roots from leaf axils; sieve tube plastids also with peripheral fibers; leaves M-shaped when mature, spiny; inflorescence bracts usually colored. - 3/805. W. Africa to the Pacific.

• Pandanaceae are woody plants with long, spiny-toothed leaves that are curved-flat in bud; the inflorescence is terminal and usually (elongated-)capitate and the perianth is at most vestigial.

CYCLANTHACEAE A. Richard - Leaves petiolate, basically simple, plicate or variants, often divided deeply; inflorescence a congested spike [spadix]; ovules tenuinucellate, micropyle bistomal. - 12/225: Central and tropical South America.

Carludovicioideae - T 10-30, with abaxial gland; inflorescence with spathe, flowers in spirals; staminate flowers: T in one or two whorls, 6+, filaments swollen basally; carpellate flowers: 4(-8)-merous, staminodes opposite P, long-filiform; fruit baccate. - 11/225. Central and tropical South America.

Cyclanthoideae - Subepidermal sclereids +; non-articulated laticifers +; lysigenous air spaces with transverse septae +; inflorescence with whorls of staminate and carpellate "flowers"; staminate flowers: A in 4 rows per whorl, connate basally; carpellate flowers: ovary cavity with closely-set placentae; fruit dry, syncarpous, splitting down the middle; seeds embedded in mucilage. - 1/1. Central and N. South America, the Lesser Antilles.

• Cyclanthaceae are more or less herbaceous and have large, petiolate, plicate, toothed and/or deeply-divided leaves. The inflorescence is spadiciform and the flowers have a more or less inferior ovary. Cyclanthus itself can be recognised by its bifid leaves (or there is a bifid midrib) and distinctive spadix with spirally-arranged flowers. Although Cyclanthaceae can be confused with palms, their petioles are usually notably softer than those of palms and may be rounded.

LILIALES + ASPARAGALES + COMMELINIDS: ?

LILIALES Perleb  

Plants rhizomatous geophytes; sleaves elliptical, fine venation reticulate, base not sheathing; T large, 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 mycotrophic herbs; leaf sheath closed; flowers single, monosymmetric, 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.

• Corsiaceae are echlorophyllous mycotrophic herbs that have leaves with closed sheaths and large, single, terminal monosymmetric flowers that are held in the "inverted" condition; the mediam member of the outer whorl is much larger than all other tepals and is adaxial. The six stamens are extrose and the ovary is inferior.

CAMPYNEMATACEAE Dumortier - Ovules crassinucellate. - 2/4. New Caledonia and Tasmania.

Petermanniaceae + Alstroemeriaceae + Luzuriagaceae + Colchicaceae + Melanthiaceae + Philesiaceae + Smilacaceae + Rhipogonaceae + Liliaceae: ?

Petermanniaceae + Alstroemeriaceae + Luzuriagaceae + Colchicaceae: 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. - 1/1. Central part of the E coast of Australia.

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

Colchicaceae + Alstroemeriaceae + Luzuriagaceae: ?

COLCHICACEAE de Candolle - 19/225. Temperate to tropical, but not in South America.

• Colchicaceae are cormose or rhizomatous geophytes that may be recognised by their rather large flowers with six free tepals, each more or less enveloping a stamen, and a superior ovary.

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.

• Alstroemeriaceae may be recognised by their spiral, leaves that are twisted at the base and so are more or less "upside-down" and have quite broad blades, their large, monosymmetric flowers which are held with the median member of the outer whorl adaxial, and their inferior ovary.

LUZURIAGACEAE Lotsy - Stems perennial; leaves 2-ranked, resupinate. - 2/5. Peru to Tierra del Fuego, Falkland Islands, New Zealand and Australia (New South Wales to Tasmania).

• Luzuriagaceae are climbing monocots with more or less woody stems that may be recognised by their two-rankedleaves that are twisted at the base and so are "upside-down" and white polysymmetric flowers with plain-colored tepals and superior ovary.

MELANTHIACEAE Batsch - 16/170. N. temperate, esp. East Asian 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: ?

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

PHILESIACEAE Dumortier - 2/2. Chile.

RHIPOGONACEAE Conran & Clifford - Stem prickly; leaves opposite; tapetum amoeboid. - 1/6. New Zealand to New Guinea.

• Rhipogonaceae are climbers that often have prickly stems. The leaves are often opposite, petiolate, and with a blade that has three, strong longitudinal veins and reticulate fine venation. The flowers are rather small and undistinguished; the fruit is a berry.

Smilacaceae + Liliaceae: ?

SMILACACEAE Ventenat - Stem prickly; leaves with lateral ligules and/or thorns or tendrils, with petiole, lamina, midrib; inflorescence umbellate; pollen inaperturate, spinulose. - 2/315. Pantropical to temperate.

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

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.

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

ASPARAGALES + COMMELINIDS: ?

ASPARAGALES Bromhead

Chelidonic acid; inflorescence racemose, 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 mycotrophic; 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. - Tropical.

• Orchidaceae can be recognised by their strongly monosymmetric flowers in which the 1-3 stamens are at least basally adnate to the style. The flowers are usually held with the median member of the outer tepalline whorl in the adaxial position, while the median member of the inner whorl is often remarkably elaborated and forms a labellum. The ovary is inferior and the fruit, a capsule, opens down the sides to release the very numerous and minute seeds.

• Apostasioideae have 2-3 stamens that are connate only to the base of the style and apiculate, carinate tepals. The other subfamilies have a well developed labellum and the stamen(s) are more or less completely fused with the style, forming a gynostemium.

• Cypripedioideae have two stamens and a deeply saccate labellum. The other subfamilies have but a single stamen. In the other subfamilies the labellum is very variable in shape, but not often deeply saccate. Vanilloideae lack both pollinia and viscidia, while both are present in Epidendroideae and Orchidoideae. The former have rather tough leaves and stems and incumbent anthers, the latter, softer, deciduous leaves and straight anthers.

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 (Baker) M. W. Chase, Rudall & Conran - 2/12. Australia.

• Boryaceae are xeromorphic, some being resurrection plants, with aerial stems and stilt roots; the inflorescence is scapose and involucrate, but the flowers are undistinguished.

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.

• Lanariaceae are monocots that can be recognised by their shortly branched inflorescences covered by dendritic hairs and radially symmetric flowers with tepals connate half-way.

Asteliaceae + Hypoxidaceae: rosette-forming or caespitose; flavonols +; mucilage canals +; endosperm thin-walled; cotyledon not photosynthetic, ligule long.

ASTELIACEAE Dumortier - 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; tapetum plasmodial; seed with prominent raphe. - 7-9/100-220. Seasonal tropics, esp. southern Africa (temperate).

• Hypoxidaceae can be recognised by their rosettes of plicate or at least folded leaves and persistent leaf bases; non-glandular indumentum is quite prominent. In the flowers, the outer tepal whorl tends to be green outside, and the ovary is inferior; there is often a narrowly tubular part of the tepal or an apical beak to the ovary.

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).

• Tecophilaeaceae are cormose herbs sometimes with broad-bladed, petiolate leaves; their flowers have spreading tepals and are monosymmetric because of the androecium, the stamens being strongly dimorphic. The anthers open by pores.

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

DORYANTHACEAE R. Dahlgren & Clifford - Pollen trichotomosulcate. - 1/2. E. Australia.

• Doryanthaceae are massive, tufted, perennial monocots, the leaves having entire margins but disintegrating into fibres at the apex. Their flowers are large, radially symmetrical, and with an inferior ovary.

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

IRIDACEAE Jussieu - Styloids +; leaves equitant, isobifacial; inflorescence a riphidium; T ± free, A 3, extrose, stigmas at the edges of the complex/expanded style. - 67/1800. World-wide.

1. Isophysidoideae Takhtajan - G - 1/1. Tasmania.

Crocoideae + Iridoideae: xanthone [mangiferin] +; inflorescence a riphidium; G inferior; endosperm usually nuclear.

2. Crocoideae G. T. Burnett - T connate. - 33/1033. Overwhelmingly southern African, to Europe, Madagascar and Central Asia.

3. Iridoideae - T nectaries, oil glands or oil hairs + [or most of the clade]. - 30/765. Worldwide, but esp. the spine of Central and South America.

• Iridaceae may be recognised by their usually ensiform and equitant isobifacial leaves and their large flowers with showy tepals, three extrorse stamens, complex stigma, and inferior ovary.

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.

• Xeronemataceae are large monocot herbs that may be recognised by their equitant isobifacial leaves and congested inflorescences with rather large, radially symmetrical, upwardly-facing flowers. The stamens are strongly exserted.

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.

• Agapanthaceae are rather robust, rhizomatous herbs that can be recognised by their rather fleshy and 2-ranked leaves and their scapose umbellate inflorescence of generally large flowers with superior ovaries.

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.

• Alliaceae can be recognised by their smell, their often rather fleshy and soft leaves, and their scapose umbellate inflorescence with rather small flowers that have superior ovaries.

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.

• Amaryllidaceae are usually bulbous herbs that can be recognised by their rather fleshy and 2-ranked leaves and their scapose umbellate inflorescence of generally large flowers with an inferior ovary.

Laxmanniaceae + Asparagaceae + Ruscaceae + Themidaceae + Hyacinthaceae + Aphyllanthaceae + Agavaceae [= Asparagaceae s.l.]: ?

A rather well supported clade in molecular analyses, but with no obvious synapomorphies, although "endosperm helobial, thick-walled, pitted, hemicellulosic" may be placed at this level. The families are difficult to recognise, for the most part having rather small and undistinguished "lily"-type flowers, hence using Asparagaceae s.l. for the clade may well be helpful.

Aphyllanthaceae + Themidaceae + Hyacinthaceae + Agavaceae:

Aphyllanthaceae + Themidaceae + Hyacinthaceae + Agavaceae: ?

APHYLLANTHACEAE Burnett - Stems photoysnthetic; leaves as non-photosynthetic scales; inflorescence scapose, flowers multibracteolate; T marcescent, pollen spiraperturate, 1 ovule/carpel. - 1/1. W. Mediterranean.

• Aphyllanthaceae may be readily recognised; the above-ground plant consists of tufts of scapose infloresences, the scapes being the main photosynthetic organs since the scarious leaves at their bases are non-photosynthetic; the inflorescence is few-flowered, the spreading, usually blue tepals being moderate in size (the whole plant looks rather like Sisyrinchium!).

Themidaceae + Hyacinthaceae: leaves spiral; pedicels bracteate; cotyledon not photosynthetic.

THEMIDACEAE Salisbury - 12/62. S.W. North America, to British Columbia and Guatemala.

• Themidacaeae are like Alliaceae in their scapose, umbellate inflorescence, rather small flowers, more or less connate tepals sometimes with a corona, and a superior ovary, but the plants have a fibrous corm, not a bulb, and lack the distinctive smell common in the latter family, and there are usually 4 or more inflorescence bracts that are not as all-enveloping as the ca 3 bracts of Alliaceae.

HYACINTHACEAE Borkhausen - 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 America.

Asparagaceae + Ruscaceae: flowers small?

ASPARAGACEAE Jussieu - Embryo sac curved, nucellar epidermal cells enlarged. - 2/165-295. Old World, but not the Antipodes (except N. Australia), Mexico.

RUSCACEAE Sprengel - 26/475. N. hemisphere, esp. Southeast Asia-Malesia (Convallariaceae s. str.), Europe and the Near East (Ruscaceae s. str.), S.W. North America (Nolinaceae s. str.), Africa, esp. the Cape and S.W. (Eriospermaceae s. str.).

COMMELINIDS

Unlignified cells walls with UV-fluorescent ferulic and coumaric acids; SiO₂ bodies in leaves; P fully bicyclic [A adnate to C/inner P], pollen starchy.

Unplaced:

DASYPOGONACEAE Dumortier - 4/16. Southern Australia.

ARECALES Bromhead  

Plant woody, usu. monopodial; cuticular waxes as aggregated rodlets, stomata tetracytic; leaves spiral, petiolate, reduplicate-plicate, pinnately pseudocompound; flowers ± sessile, 1 apotropous ovule/carpel; embryo short, broad. - 1 family, 187 genera, 2000 species.

ARECACEAE Schultz-Schultzenstein//Palmae Jussieu - 190/2000. Humid tropics and subtropics (warm temperate).

1. Calamoideae Beilschmied - Fruits with recurved scales. - 22/. Tropical, but esp. Sri Lanka to West Samoa and Fiji.

2. Nypoideae Griffith - Branching dichotomous; leaves reduplicate; inflorescence with axes adnate to internode above; carpels separate. - 1/1. Malesia (Bengal to Queensland).

3. Coryphoideae Burnett - Leaves induplicate. - 44/ . Africa, esp. South East Asia.

4. Ceroxyloideae Drude

5. Arecoideae Burnett - 112/ . Pantropical.

• Arecaceae can be recognised by their woody stems, large, tough, petiolate leaves with plicate and often apparently compound blades, axillary inflorescences (but Corypha, etc.!) with numerous flowers, and 1-seeded fruits. Cyclanthaceae can be similar vegetatively, but their petioles are usually less woody and the leaf never has a hastula, an adaxial flap-like structure at the apex of the petiole.

POALES + COMMELINALES + ZINGIBERALES: Primary cell wall mostly with glucurono-arabinoxylans; stomata paracytic or tetracytic with parallel cell divisions; endosperm starchy.

POALES Small  

Vessels also in stem and leaf; SiO₂ epidermal; raphides 0; endosperm nuclear, embryo broad, 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 - 1/14. Temperate and Arctic, to New Zealand.

• Sparganiaceae are rhizomatous aquatics with linear leaves and inflorescences with a few spherical heads of flowers, carpellate towards the base of the inflorescence and staminate towards the apex.

TYPHACEAE Jussieu - 1/8-13. Temperate and tropical regions worldwide.

• Typhaceae are rhizomatous herbs that like damp conditions and have erect, linear leaves and distinctive, elongated, densely cylindrical inflorescences, the part containing the carpellate flowers being borne below and separate from the part with the staminate flowers.

BROMELIACEAE Jussieu - Silica bodies solitary; hairs water-absorbing; leaves lacking a sheath; 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.

• Bromeliaceae are often rosette plants, the leaves always being spirally arranged, lacking a distinct sheath, and often with spiny-toothed margins; the indumentum is lepidote; and the terminal inflorescence is prominently bracteate. The flowers have a calyx and corolla, and the latter, although free, is erect and forms a tube.

Rapateaceae + Cyperaceae group + Poaceae group: ?

RAPATEACEAE Dumortier - Inflorescence capitate, scapose; 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.

• Rapateaceae are rosette herbs with capitate, scapose inflorescences bearing rather conspicuous flowers; the perianth is differentiated into calyx and corolla, there are six stamens, and the anthers dehisce by pores.

Rapateaceae + 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.

• Thurniaceae can be recognised by their erect stems, serrate leaves, and pentacyclic flowers with scarious perianth.

Juncaceae + Cyperaceae: micropyle endostomal.

JUNCACEAE Jussieu - Stems round. - 7/430. Worldwide, esp. Andes (3 endemic genera), S. South America-New Zealand (2 endemic genera).

• Juncaceae are glabrous (Luzula excepted) caespitose herbs that can be recognised by their solid, rounded stems, leaves that are also often rounded and have open sheaths, and pentacyclic flowers with a scarious perianth.

CYPERACEAE Jussieu - SiO₂ bodies conical, attached to walls; 1 basal ovule/flower. - 98/4350. World-wide.

• Cyperaceae are herbs with solid, sharply three-angled stems and leaves with closed sheaths. The flowers are very reduced and often aggregated into spikes or heads, the perianth, when recognisable, is scarious, and the fruit is an achene.

Mayacaceae + Eriocaulaceae + Xyridaceae + Anarthriaceae + Restionaceae + Centrolepidaceae + Flagellariaceae + Joinvilleaceae + Ecdeiocoleaceae + Poaceae: ovules tenuinucellate; embryo minute.

Mayacaceae + Eriocaulaceae + Xyridaceae: flavonoids +; SiO₂ bodies 0; leaves spiral; inflorescence axillary, scapose, capitate, with involucral bracts; flowers moderate in size; P = K + C, A 3, basifixed, pollen more or less spiny, style +; K persistent in fruit; seed operculum ["embryostega"] +, endostomal; embryo undifferentiated.

MAYACACEAE> Kunth - Leaves flat, apically bidentate, univeined, without a distinct sheath, uniseriate colleters +; flower single, apparently axillary; ovules atropous, micropyle bistomal. - 1/4-10. Mostly tropical and American (inc. S.E. U.S.A.), 1 sp. from W. Africa.

• Mayacaceae are small herbs of marshes looking rather like a club-moss. They have numerous spirally-arranged, apically-toothed leaves borne scattered along the stem and pink to white apparently axillary flowers with a clearly differentiated calyx and corolla.

Eriocaulaceae + Xyridaceae: rosette plants; vessel elements with simple perforations; C clawed, A adnate and opposite C, exothecium +.

ERIOCAULACEAE Martynov - Flowers imperfect; inflorescence receptacle ± flat; pollen spiraperturate, 1 pendulous atropous ovule/carpel. - 10/1160. Pantropical (to temperate).

1. Eriocauloideae - Plants usu. of aquatic habitst; 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.

• Eriocaulaceae are nearly always rosette plants readily recognised by their “pipe-brush” inflorescences. These have slender peduncles that are more or less spirally twisted and ridged or angled, and many small flowers with rather scarious and inconspicuous corollas that are aggregated into dense heads. The base of the inflorescence is surrounded by a single closed leaf sheath.

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; staminodia 3, branched and with moniliform hairs on branch ends, placentation (intrusive) parietal. - 1/225-300. Pantropical to warm temperate, esp. Brasil.

2. Abolboidoideae - Pollen spherical, inaperturate, style solid. - 4/26. W. South America, Guyana Highlands in particular.

• Xyridaceae are rosette plants that sometimes have equitant leaves; they can be recognised best by their inflorescences and flowers. The inflorescences normally bear one or more pairs of bracts below the flowers. The latter are borne in heads and have a conspicuous but ephemeral corolla.

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 - 3/11. West Australia.

Centrolepidaceae + Restionaceae: anthers bisporangiate/unithecal, embryo sac with compound starch grains, cells of nucellar epidermis anticlinally elongated.

CENTROLEPIDACEAE Endlicher - 3/35. Hainan, IndoChina and Malesia to New Zealand, S. South America (Gaimardia).

• Centrolepidaceae are rather small more or less caespitose herbs. The inflorescence is scapose and spicate or capitate and involucrate.

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 (Apodasmia).

• Restionaceae are sometimes quite large herbs with very much reduced leaves; the flowers are small, imperfect, but usually with a perianth, and aggregated into spikelets.

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; leaf blade terminating in a tendril. - 1/4. Paleotropics, to the Pacific Islands.

• Flagellariaceae are robust, climbing or scrambling rather grass-like plants that can be recognised by their leaves that terminate in a sensitive tendril by which the plant is attached to its support. The rather large, paniculate inflorescences have many small, perfect flowers.

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 Tomlinson & A. C. Smith - Flowrs perfect. - 1/2. Malay Peninsula to the Pacific.

• Joinvilleaceae are robust, grass-like plants with plicate leaves and terminal panicles of small, perfect, brownish flowers.

Ecdeiocoleaceae + Poaceae: 2 adaxial tepals of the outer whorl larger than the rest; pollen with operculum, wall without scrobiculi, with intraexinous channels.

ECDEIOCOLEACEAE D. F. Cutler & Airy Shaw - Leaves lacking a blade. - 2/2. W. Australia.

POACEAE (R. Brown) Barnhart - Sieve tube plastids also with rod-shaped protein bodies; leaves pseudopetiolate, midrib +, ligulate; P ?, G with 1 central ovule; fruit an achene, the testa closely adherent to pericarp, hilum long; peripheral layer of endosperm meristematic, embryo lateral, long, well differentiated, plumule lateral, cotyledon = absorbtive scutellum; collar [epiblast, the ligule of the cotyledon] conspicuous; trnT inversion in chloroplast genome. - 668/10025. Worldwide.

1. Anomochlooideae Potzdal - 2/4. Brasil, forests.

Pharoideae + Puelioideae + PACCAD + 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 and palea [?= connate adaxial tepals of the outer whorl]; lodicules [inner whorl tepals] 3; n = 12.

2. Pharoideae (Stapf) L. G. Clark & Judziewicz - Microhairs 0; leaves resupinate, lateral veins oblique. - 2/12. Pantropical, in forests.

Puelioideae + PACCAD + 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.

PACCAD + BEP clades: arm and fusoid cells 0; foliar cross veins 0; pseudopetiole 0; A 3, lodicules 2, stylodia free. Mostly non-forest.

Panicoideae + Arundinoideae + Centothecoideae + Chloridoideae + Aristidoideae + Danthonioideae [PACCAD clade]: mesocotyl internode elongated, epiblast 0.

Panicoideae + Centothecoideae: hilum non-linear.

4. Panicoideae Link - 206/3270. Tropics to temperate.

5. Centothecoideae Soderstrom - /45. Warm temperate to tropical forests. Poorly understood.

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 - /1400. Tropical to warm temperate, more or less dry environments especially in Africa and Australia.

Aristidoideae + Danthonioideae: awn trifid, or 3 awns.

8. Aristidoideae Caro - 3/385. Warm temperate, few in Europe.

9. Danthonioideae Barker & Linder - 19/250. Mesic to xeric.

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

10. Bambusoideae Luersson - /1200. Tropical, often in forests.

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

12. Pooideae Bentham - /3300. Largely N. temperate.

• Poaceae can be recognised by their leaves that have long sheaths and ligules at the sheath-lamina junction, round and often hollow stems, and inflorescences whose basic unit is a spike. Individual flowers are small, lacking an obvious perianth and with a gynoecium that usually has two plumose stigmas and a single ovule. In the dry, achenial fruit (often called a caryopsis), the rather large embryo is lateral, lying next to the seed coat on one side.

COMMELINALES + ZINGIBERALES: inflorescences with many-flowered cincinnal [helicoid cyme] branches; tapetum invasive or plasmodial.

COMMELINALES Dumortier  

Vessel elements scalariform; seed coat testal and tegmic; endosperm abundant, helobial, cell wall formation in small chalazal chamber precedes that in large micropylar chamber. - 5 families, 68 genera, 812 species.

Commelinaceae + Hanguanaceae: cotyledon not photosynthetic.

COMMELINACEAE Mirbel - 40/652. Tropical and temperate.

1. Cartonematoideae (Pichon) G. Tucker - SiO₂ bodies 0; one flower/cincinnus; 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 +. - 38/640. Tropical, also temperate, not Europe.

• Commelinaceae are herbs with rather soft and fleshy leaves; these have closed cheaths and may be involute in bud. The inflorescences usually consist of many-flowered helicoid cymes that are often subtended by large, leafy bracts; they are quite often leaf-opposed. The flowers usually last for one day only and are frequently blue or pink; when they are monosymmetric they are usually held with the median petal in the abaxial position. The filaments are often conspicuously hairy.

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.

• Hanguanaceae are robust terrestrial herbs that have petiolate leaves with a midrib, many parallel veins, and prominent cross veins. The inflorescence is a panicle with sessile, small, and undistinguished flowers. The fruit is a single-seeded berry, and the seed itself is bowl-shaped as it comes to surround the placenta during development.

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. - 4/5. Australia (all genera) to Southeast Asia.

• Philydraceae can be recognised by their 2-ranked, ensiform leaves - although Philydrella lacks these vegetative characters - inflorescences with wooly hairs and distinctive monosymmetric flowers. These have a two-merous perianth, the outer whorl usually being much larger than the inner and a single stamen.

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.

Conostyloideae Lindley - Hairs branched; flowers enantiostylous, pollen porate, placentae with tanniniferous cells; cotyledon photosynthetic, hypocotyl +, primary root well developed. - 6/77. S.W. Australia.

• Haemodoraceae may be recognised by their 2-ranked, equitant-isobifacial leaves, their inflorescences, which are often rather elongated bifurcated helicoid cymes, their sometimes monosymmetric and/or enantiostylous flowers with inferior ovaries, and their roots, which in Haemodoroideae have a distinctive red color.

PONTEDERIACEAE Kunth - Filaments hairy, pollen 2- or 3-sulcate. - 9/33. Tropics, also temperate, esp. New World.

• Pontederiaceae are aquatic or marsh plants - sometimes annuals - with sheathing leaf bases that are sometimes differentiated into a petiole and blade with many parallel veins, the blade of the younger leaf then encircles the petiole of the older leaf. The inflorescence has two bracts, either/both leaf-like and/or sheathing, and the marcescent, monosymmetric flowers are either basically blue or yellow.

ZINGIBERALES Grisebach  

No aerial stem except when flowering; SiO₂ in bundle sheath; sieve tube plastids also with starch grains; petiole bundles in arcs; cuticular waxes as aggregated rodlets; leaves with distinct petiole, midrib and fine cross venation; inflorescence bracts large, persistent; flowers large, monosymmetric; T = colored K surrounding C, A 5, adaxial member of inner whorl not developed, anthers long, pollen inaperturate, exine thin and spinulose to 0 [pollen not resistant to acetolysis], G inferior, outer integument 3< cells across, nucellar epidermal cells radially elongated, style long, stigma large, wet; fruit capsular; seeds arillate, operculate, operculum testal, micropylar collar [develops from outer integument and forms annular inpushing in perisperm surrounding operculum], endotesta sclerotised and silicified; endosperm nuclear, perisperm s.l. +, starchy, embryo plug-like; cotyledon not photosynthetic, ligulate, collar roots +. - 8 families, 92 genera, 2111 species.

MUSACEAE Jussieu - Bracteoles. - 2/35. Africa, Himalayas to South East Asia, Philippines and N. Australia.

• Musaceae are large herbs with spiral, shortly-petiolate leaves in which the secondary veins depart more or less at right angles from the midrib. The inflorescence bracts are usually deciduous and subtend fascicles of ebracteate flowers. The monosymmetric flowers have five tepals connate except adaxially, where there is a single, free, deeply concave tepal.

HELICONIACEAE Nakai - Median outer T adaxial, free, 5 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.

• Heliconiaceae are large herbs with 2-ranked leaves and inflorescences with large, colored bracts in the axils of which are fascicles of flowers with petaloid tepals; the drupaceous fruit is borne on a stout, elongated pedicel and has a single hard stone per loculus.

Strelitziaceae + Lowiaceae: petiole with adaxial and abaxial series of air canals; adaxial A of inner whorl sterile, tapetum glandular [inc. Lowiaceae?], floral column [sterile apex of ovary] +, stigma 3-lobed; aril hairy.

STRELITZIACEAE Hutchinson - 3/7. Tropical South America, E. southern Africa, Madagascar.

• Strelitziaceae can be recognised by their 2-ranked, long-petiolate leaves and inflorescences with tough, boat-shaped, green inflorescence bracts. The distinctive flowers have more or less connate lateral abaxial petals, and the woody capsules have seeds with brightly colored, fimbriate arils.

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, adaxial C large, labellar, abaxial 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.

• The leaves of Lowiaceae have steeply ascending and apparently rather distant longitudinal veins (some are much more prominent than the others) and strongly developed cross veins that are more evident on the abaxial than the adaxial surface. The flowers are held with the median sepal adaxial, the median petal forms a well-developed abaxial labellum, and the two lateral petals are small.

Cannaceae + Marantaceae + Zingiberaceae + Costaceae: raphides 0; petiole with one series of air canals; guard cells with inner and outer ledges unequal; petiole short, poorly differentiated; C connate, both A whorls with two staminodes, adaxial A of inner whorl fertile; micropylar collar well developed, cells of exotesta longitudinally elongated; endosperm slight, chalazosperm +.

Cannaceae + Marantaceae: oblique cells in petiole; flowers asymmetrical; A ½, staminodes free, stigma not notably expanded.

CANNACEAE Jussieu - Mucilage canals in stem; style flattened; capsule muricate. - 1/19. New World (sub)tropics.

• Cannaceae are large herbs with petiolate leaves that may be recognised by their large, short-lived, asymmetrical flowers with a flattened style and muricate inferior ovary; operculum 0, mesotesta sclereidal, endotesta not sclereidal.

MARANTACEAE R. Brown - Petiole well differentiated; flowers explosive. - 31/550. Tropics, esp. American, not in Australia.

• Marantaceae are often robust herbs that may be recognised by their pulvinate, petiolate leaves and paired, often only moderately-sized, asymmetrical flowers. The vegetative plant is made up of repeating units with prophyll, reduced leaf, and then leaves varying in number and internode length - and also orientation, since the plane of distichy may be at right angles to that of the parent axis.

Zingiberaceae + Costaceae: leaf ligulate; K connate, labellum from lateral staminodes of outer whorl and from the 2 staminodes of inner whorl, large, with narrow tube and distinct open limb, A 1, exine +, style slender, running between two half anthers, stigma cup- or funnel-shaped; chalazal mass in seed ± developed, endosperm helobial; hypocotyl well developed.

ZINGIBERACEAE Martynov - 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; 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.

• Zingiberaceae are usually large and often aromatic rhizomatous herbs with 2-ranked leaves that have open sheaths; an inflorescence that is frequently branched; and pedicellate, monosymmetric flowers with a single stamen, the style running between the two halves of the large anther.

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.

• Costaceae can be recognised even vegetatively: they have ligulate leaves with a closed sheath that are arranged in a single spiral up the stem. Their inflorescences are dense, spicate-capitate, and have large bracts, and their monosymmetric flowers have a large labellum and single stamen, the style running between the two halves of the large anther.

EUDICOTS: Ethereal oils 0; 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?

RANUNCULALES Dumortier

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; leaves spiral, 2ndary veins palmate; seed exotestal; endosperm development?, embryo size? - 7 families, 199 genera, 4445 species.

EUPTELEACEAE K. Wilhelm - Carpels stipitate, stigma brush-like; fruit samaroid. - 1/2. Temperate South East Asia.

• Eupteleaceae can be recognised by their spiral, dentate leaves with strong, ascending, pinnate veins going into the teeth, the excavated petiole bases, and the small flowers that lack a perianth and have anthers with a long connective and separate stipitate carpels. These latter develop into small, disc-shaped samaras down one side of which are the remains of the brush-like stigma.

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

Lardizabalaceae + Circeasteraceae + 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.

Lardizabalaceae + Circaeasteraceae: A extrorse.

LARDIZABALACEAE R. Brown - Flowers imperfect. - 9/36. South East Asia and Chile.

• Lardizabalaceae are viny, monoecious or dioecious plants with compound leaves and broad rays in the wood. The medium-sized, three-merous flowers have the perianth members and androecium opposite each other. The carpels are free and the fruits are more or less fleshy, although they are sometimes follicles.

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

• Circaeasteraceae are easily recognised. They are small herbs, their leaves have largely dichotomous venation, and their rather small flowers have separate carpels.

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

MENISPERMACEAE Jussieu - Lianes; petiole pulvinate at apex and base; flowers imperfect, P with a single trace, pollen with endoapertures [colporate], ovules 2/carpel, 1 aborting; fruit a drupe. - 70/420. Pantropical, usually lowland.

• Menispermaceae are recognisable by their often viney/lianescent habit, broad rays, petioles pulvinate at both ends, and often coriaceous lamina with palmate venation. The lamina is also often peltate, and even if not, the two sides of the lamina nearly meet on top of the petiole. The plants are dioecious, the often 3-merous flowers are small, and the drupelets often have a distinctively sculpted endocarp and can be strongly curved (hence "moon-seed").

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 +); nucellar cap +, outer integument at least 4 cells thick; seed exotestal.

BERBERIDACEAE Jussieu - G 1. - 14/701: mostly East Asia and E. North America, also South America, N. Africa, general N. temperate.

1. Berberidoideae - 13/700. - Mostly East Asia and E. North America, general N. temperate, also South America, N. Africa; many genera disjunct.

2. Nandinoideae - K many, spiral, nectary 0. - 1/1. East Asia.

• Berberidaceae are herbs or shrubs that may be recognised by their often yellow-colored secondary tissue (most obvious in the roots) and their often compound, stipulate leaves with palmate venation and sharply toothed or spiny margins. Their flowers are usually 3-merous, the perianth is often multiseriate, the stamens are opposite the perianth, and anther dehiscence is valvate; there is always a single carpel.

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 (Rafinesque) Martynov - Petiole base on rhizome encircling stem; P members with a single trace, pollen tectum striate-reticulate, micropyle zig-zag; exotesta strongly palisade, exotegmen lignified, both multiplicative. - 1/1. C. and E. North America.

2. Glaucidioideae (Tamura) 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. - 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 - 3/17. East Asia, E. and W. North America.

Isopyroideae + Ranunculoideae: ovules apotropous [when single].

4. Isopyroideae Rafinesque - 9/450. N. temperate, also South America, Africa and New Guinea.

5. Ranunculoideae Arnott - 46/2025. Worldwide, but not tropical.

• Ranunculaceae can be recognised by their usually herbaceous habit and spiral, usually exstipulate, lobed leaves with broad bases. The flowers have large and often petaloid sepals, nectariferous "petals", spiral, usually numerous stamens, and separate carpels borne on a well-developed receptacle.

PAPAVERACEAE Jussieu - Herbaceous; roots diarch [lateral roots 4-ranked]; leaf base broad; K 2, median, distinct from C, A 4-6, G [2], placentation parietal, style +, short; capsule septicidal, (with persistent woody placenta); endotesta well developed. - 41/760. Largely N. Temperate, also S. Africa, scattered in South America, etc.

1. Pteridophylloideae - Venation pinnate. - 1/1. Japan.

Papaveroideae + Fumarioideae: 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 - Fruit dehiscing explosively from the base. - 3/16. W. North America.

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

3. Fumarioideae Eaton - Flowers strongly disymmetrical. - 17/530. Mostly N. temperate, also S. Africa.

3A. Hypecoeae Dumortier - Inner C 3-lobed, A 4, two with two vascular strands, extrorse, pollen deposited on inner petals. - 1/18. Mediterranean to W. China .

3B. Fumarieae Dumortier - 16/510. - Flowers spurred, inner C connate, A in two groups of three, 2 anthers dithecal and 4 monothecal, pollen deposited on stigma. - N. temperate, especially Mediterranean, inc. N. Africa (S. Africa).

• Papaveraceae are usually herbs whose leaves have broad bases. The flowers are 2-merous, and the calyx and corolla are clearly distinguishable; the gynoecium has two (or more) carpels, parietal placentation, and usually a short style.

• Pteridophylloideae are herbs with a rosette of fern-like leaves and a scapose inflorescence with rather small, 2-merous flowers which have a distinct style.

• Papaveroideae are usually herbs that may be recognised by their often soft if rather bristly-hairy leaves, copious latex, and flowers with two, large, fugaceous sepals enclosing the bud, crumpled petals, usually numerous stamens, and syncarpous gynoecium with at most a short style and parietal placentation.

• Fumarioideae are often fleshy herbs with watery sap and distinctive 2-merous mono- or disymmetrical flowers that are usually spurred; the calyx is very small or almost invisible.

SABIALES + PROTEALES + TROCHODENDRALES + BUXALES + GUNNERALES + CORE EUDICOTS: (axial nectary +).

SABIALES Takhtajan  

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

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

• Sabiaceae are evergreen trees or lianes that may be recognised by the brochidodromous venation of their leaves or leaflets, their small flowers with the perianth whorls and stamens all opposite, and the more or less flattened and distinctly curved drupes; the surface of the latter is often distinctly venose.

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.

• Nelumbonaceae are water plants that are easily recognised by their peltate leaves with dichotomous main venation that are held above the water surface like parasols, and their flowers which are like those of water lilies but have free carpels immersed in a large, obconical receptacle.

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.

• The bark exfoliates with characteristic plates, the leaves have leaf-like stipules completely surrounding the stem, the petiole base encloses the axillary bud, and the blades have subpalmate venation. However, Platanus kerrii is rather different vegetatively from the other species, having stipules that are more or less scarious, an axillary bud that is not enclosed by the petiole base, and pinnate venation. In all species, the inflorescences are capitate, the staminate flowers have valvate anthers and the carpellate flowers have free carpels; the fruits are small achenes with tufts of basal hairs.

PROTEACEAE Jussieu - Hairs with 2 short cells, one in epidermis, apical cell elongated; stipules 0; bracteoles 0; P decussate, pollen triangular in polar view, pores broadly operculate, 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 + Proteoideae + Grevilleoideae: P connate, A adnate to P, more or less sessile, nectaries +.

2. Persoonioideae - Chromosomes 13-26 µm long. - 5/110. Mostly Australia, also New Caledonia and New Zealand.

Proteoideae + Grevilleoideae: flowers vertically or obliquely monosymmetric, P split to base on one side, (secondary pollen presentation, the apex of the style bearing the pollen).

3. Proteoideae - Al accumulation infrequent; flowers sessile; fruit single-seeded. - 26/640. S. Africa, Australia.

4. Grevilleoideae Engler - Al accumulation +; paired flowers +. - 45/855. Australia, to Southeast Asia, South America, Africa and Madagascar.

• The family may be recognised by the broad rays in the wood, the surface of which is often furrowed, the often rather coriaceous leaf blades with rather distant (or no) teeth and prominent reticulum, the swollen leaf base and the sometimes rather short petiole. The 4-merous flowers are often aggregated into conspicuous inflorescences; the perianth may be monosymmetric and curved and the stamens are opposite the perianth members; the single carpel often has a long style.

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

TROCHODENDRALES Cronquist  

Evergreen trees; vessels 0; 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.

• Trochodendraceae are evergreen trees may be recognised by their serrate, exstipulate leaves with more or less palmate venation and their flowers in which there are several laterally connate carpels arranged in a whorl. The stylodia become basal in the follicular fruits.

BUXALES + GUNNERALES + 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. - 1/2. E. Madagascar.

• Didymelaceae may be recognised by their spiral, entire, greenish-drying leaves, indumentum of small scales, and small flowers, the staminate flowers having two extrorse anthers and the carpellate flowers a single carpel with a long, oblique stigma.

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

1. Sarcococceae - 3/22. E. Asia to W. Malesia, E. North America, N. South America.

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

• Buxaceae are generally monoecious evergreen herbs to small trees with exstipulate and often opposite leaves and rather small flowers with an inconspicuous perianth. The stamens are opposite the perianth, and the stylodia, stigmatic their entire length, are prominent and more or less spreading. The fruit is few-seeded, the mesocarp and endocarp separate, and the latter splits septicidally as well. In flower and fruit Buxaceae look rather like Euphorbiaceae, but there is no columella in the fruit, unlike Euphorbiaceae.

GUNNERALES + 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.

GUNNERALES Reveal

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.

• Gunneraceae are sometimes huge herbs that can be recognised by their spiral leaves the blades of which have palmate venation that is very prominent on the lower surface, often rather stipule-like cataphylls, and their relatively large and branched inflorescences bearing many small, inconspicuous flowers; the ovary is inferior and bicarpellate.

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.

• Myrothamnaceae are resurrection plants; the leaves may appear to dry out, but they regain turgor, etc., when conditions improve. They are vegetatively very distinctive aromatic shrubs; the leaves are opposite and have a long, common sheath on which there are four persistent stipular points, and the blades have palmate secondary veins and toothed margins.

CORE EUDICOTS: Root apical meristem closed; flowers rather stereotyped: 5-merous, K and C distinct, K with 3 traces, A = 2x K, (often numerous, but then often 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].

K has three traces, C has one; taxa with numerous stamens, often developing from 5(10) primordia and/or centrigual, are found in most orders.

BERBERIDOPSIDALES Doweld  

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; fruit a drupe; embryo transverse. - 1/1. Chile.

• Aextoxicaceae are dioecious evergreen trees that can be recognised by their opposite, entire exstipulate leaves that are covered with peltate scales and by their pendulous, racemose inflorescence. The petioles are almost pulvinate at the apex and the base and the lamina appears to be minutely peltate. The flower buds are covered by bracteoles and when they fall off, the almost scale-like sepals fall off with them.

BERBERIDOPSIDACEAE Takhtajan - 2/3. Chile, E. Australia.

• Berberidopsidaceae are woody scramblers that can be recognised by their exstipulate, sometimes spiny-toothed leaf blades with palmate venation; the veins run straight into the spines, when present. The fruit is a berry crowned by the persistent style base.

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.

DILLENIALES Hutchinson  

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

DILLENIACEAE Salisbury - 10/300. Tropical and warm temperate.

• Dilleniaceae are recognisable by the often strong and parallel secondary veins that proceed straight into the teeth; scalariform tertiary venation is common, and the lamina is often rough; the leaves tend to elongate when they are still rolled up. The bark is often rich brown. The pedicels are articulated near the apex and persist after the flower falls off; the flowers themselves are often conspicuous and have crumpled petals and numerous stamens that are reflexed in bud and often have porose anthers. The fruits are shiny follicles containing arillate seeds; the calyx is persistent, sometimes accrescent, and the filaments are also persistent.

CARYOPHYLLALES Perleb  

Not mycorrhizal; root hair cells in vertical files; perforation plates not bordered; leaves entire; anther with outer parietal cells developing directly into the endothecium, pollen colpate, when G = K or P, opposite them, micropyle endostomal, styles/branches long; seed coat? - 31 families, 692 genera, 11155 species.

RHABDODENDRACEAE (Huber) Prance - Sieve tube plastids with protein crystalloids and starch; secretory cavities with resin; foliar fibre-like sclereids +; hairs peltate, fringed; leaves revolute, obscurely punctate; A many, filaments very short, anthers very long, G 1, with 1 (2) basal campylotropous ovules, megasporocytes several, style basal; fruit a drupe, pedicel apex swollen. - 1/3. Tropical South America.

• Rhabdodendraceae are evergreen trees that may be recognised by their fringed-peltate hairs, their rather large, entire and exstipulate leaves, flowers with stamens that have short filaments and long anthers, and a single carpel with a basal style. The fruit is distinctive, the drupes being surrounded by the persistent calyx and swollen pedicel apex. The punctations in the lamina may not be at all obvious.

(Droseraceae + Tamaricaceae + Polygonaceae groups) (Simmondsiaceae ((Asteropeiaceae + Physenaceae) (Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.))): only alternate vascular pitting; tectum spinulose; fruit a capsule.

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. - 3/115. World-wide.

• Droseraceae are insectivorous herbs usually with adaxially coiled or folded leaves and moderate-sized flowers; there are three carpels usually terminated by branched stylodia.

Nepenthaceae + Drosophyllaceae + Ancistrocladaceae + Dioncophyllaceae: rays 1-2 cells wide; cortical bundles in stem; 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; P uniseriate, with nectariferous glands adaxially, A connate into a central column, pollen in tetrads. - 1/90. Madagascar to New Caledonia.

• Nepenthaceae are insectivorous plants that are easy to recognise because of the lidded pitchers borne on the end of a twining prolongation of the leaf. The leaf base itself is broad, further widening to form a laminar portion that then narrows to form the tendrillar portion. The plants are dioecious, the inflorescences are racemes, the flowers are rather inconspicuous, and the seeds are very small.

Drosophyllaceae + Ancistrocladaceae + Dioncophyllaceae: ?

DROSOPHYLLACEAE Chrtek, Slavíkovà & Studnicka - Leaves abaxially circinate, with glandular hairs, sensitive. - 1/1. Iberian Peninsula, Morocco.

• Drosophyllaceae are subwoody carnivorous plants that may be recognised by their long, abaxially coiled leaves that have sticky, glandular hairs in long lines on their abaxial surfaces; these hairs are not sensitive. The flowers are relatively large, short-lived (they remain open only a single day), and the petals are marcescent.

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

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.

• Ancistrocladaceae are lianes that can be recognised by their shortly petiolate, extsipulate, entire leaves with glands in pits on the abaxial surface, and the complex, branched stem grapnels that are often opposite the leaves.

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; fruits open early; seeds flattened, on elongated funicles. - 3/3. Tropical West Africa.

• Dioncophyllaceae are distinctive in their rather long rosette leaves with closely parallel venation, these are followed by leaves with a short blade and glandular hairs on the abaxial surface of the prolonged midrib, and then, in the climbing phase, by leaves with paired grapnel hooks at the apex. The fruits open early and expose the still-developing flattened seeds.

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 - 5/90. Eurasia and Africa, esp. Mediterranean to Central Asia.

• Tamaricaceae are woody plants with small leaves and flowers; the styles are long and are terminated by distinctive expanded stigmas. The capsular fruits release hairy seeds.

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 an anthocarp; exotesta ± persistent, otherwise seed coat undistinguished.

PLUMBAGINACEAE Jussieu - A opposite C, ovule circinotropous, funicle long and curved, embryo sac tetrasporic, 4-nucleate [Plumbago-type]. - 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 - Deletion of rpl16 intron. - 22/800. Mostly Irano-Turanian (Mediterranean), but also S. Africa, S. South America, and W. Australia.

• Plumbaginaceae are perennial herbs or shrubs, rarely climbers. They can be recognised by their leaves which have broad bases and are frequently only shortly petiolate and/or with entire margins, and their flowers which have a relatively large, persistent, and sometimes colored calyx and antepetalous stamens. The single ovules borne at the ends of long, curved funicles are easily enough seen on dissection.

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

• The flowers of Polygonaceae are rather small, all parts (bar the carpels) are more or less free, and the 5 or 6 tepals are persistent or accrescent in fruit; the achene itself is often angled. A sheathing stipule is common and the nodes of the herbaceous taxa in particular are more or less swollen. Eriogonum and relatives lack the distinctive stipule and have a more or less cymose and involucrate inflorescence.

Simmondsiaceae ((Asteropeiaceae + Physenaceae) (Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.)): nodes 1:1; styles or style branches stigmatic their entire length; endosperm slight.

SIMMONDSIACEAE van Tieghem - Petiole articulated; staminate plant: A extrorse, anthers longer than connective, pollen ± porate, central part operculoid; carpellate plant: flowers axillary, 1 apical 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.

• Simmondsiaceae are evergreen shrubs that may be recognised by their opposite leaves that are clearly articulated at the base and have flat, rather thick blades with entire margins. The staminate flowers are small and borne in terminal inflorescences; the carpellate flowers are single and axillary and the calyx is much accrescent in fruit.

(Asteropeiaceae + Physenaceae) (Caryophyllaceae, Nyctaginaceae, Cactaceae, etc.): ?

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

ASTEROPEIACEAE (Szyszylovicz) Reveal & Hoogland - Cotyledons spirally twisted. - 1/8. Madagascar.

• Asteropeiaceae are evergreen trees that may be recognised by their shortly petiolate leaves whose blades are entire, exstipulate and have rather indistinct venation. The flowers are small are in terminal panicles, the fruit has a single seed, and the calyx is much accrescent and spreading.

PHYSENACEAE Takhtajan - Seed large, coat vascularised, 16-20 layers thick, walls not notably thickened. - 1/2. Madagascar.

• Physenaceae are rather undistinguished with their more or less 2-ranked, entire, exstipulate and coriaceous leaves. The flowers lack a corolla, the stamens have a very thin filaments and long anthers, and the styles are long. The fruit is inflated and contains a single seed which apparently has a vascularised seed coat.

Caryophyllaceae, 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; C 0, 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; 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.

• The family can be recognised by its herbaceous habit, opposite, entire leaves that often have little in the way of a petiole and are joined by a line at the base; geotropic adjustments occur at the swollen nodes. The inflorescence is cymose and the flowers appear to have a clearly distinguishable calyx and corolla, the latter being bilobed and/or clawed; there are usually 5 or 10 stamens. The fruit is a capsule.

Achatocarpaceae + Amaranthaceae: ?

ACHATOCARPACEAE Heimerl - 3/7. S.W. USA 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.

• Amaranthaceae are more or less succulent herbs often with swollen nodes. The small flowers often have 1-3 carpels and 1-2 basal ovules. The fruits either have a circumscissile capsule and a chartaceous to scarious perianth and bracts or are indehiscent and are closely invested by the more or less fleshy perianth and bracts.

Stegnospermataceae [Limeaceae [[[Corbichonia + Lophiocarpus] [Aizoaceae, Nyctaginaceae, etc.]] [Molluginaceae, Cactaceae, etc.]]: ?

STEGNOSPERMATACEAE Nakai - 1/3. Central America, the Antilles.

• Stegnospermaceae are more or less scandent or sprawling plants with rather fleshy leaves and racemose inflorescence with more or less rotate and apparently petaloid flowers.

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

[[Corbichonia + Lophiocarpus] [Aizoaceae, Nyctaginaceae, etc.]]: ?

Corbichonia + Lophiocarpus: ?

Barbeuiaceae + Aizoaceae + Sarcobataceae + Phytolaccaceae + Nyctaginaceae: ?

BARBEUIACEAE Nakai - 1/1. Madagascar.

Aizoaceae + Phytolaccaceae + Sarcobataceae + Nyctaginaceae: raphides +.

AIZOACEAE Martynov - Epidermis with bladder-like cells. - 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; capsule circumscissile; seeds arillate. - 5/36. Tropics and Subtropics.

Aizooideae + Mesembryanthemoideae + Ruschioideae: 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] and A 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 - 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.

• Aizoaceae are more or less prostrate herbs that may be recognised by their fleshy, opposite leaves the bases of which usually surround the stem and the bladder-like epidermal cells. The flowers either have five perianth members that are green outside and colored inside or numerous linear "petals". There are usually many stamens, a more or less inferior and septate ovary, and a fruit that opens on moistening as the septal tissue swells.

Sarcobataceae + Phytolaccaceae + Nyctaginaceae: ORF 2280 sequence similarity, 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.

• Phytolaccaceae can be recognised by their racemose inflorescences of apetalous flowers which have many, connate carpels - or a single carpel - each with a single basal ovule and a style; the fruit is often a berry. The plants are herbs, trees, or lianes, and the leaves tend to be rather thick and fleshy, lacking prominent fine venation.

NYCTAGINACEAE A.-L. de Jussieu - P connate, petaloid, lobes induplicate-valvate or contorted, G 1, style long; fruit achene, surrounded by accrescent (viscid) P. - 30/395. Tropical to warm temperate.

• Nyctaginaceae may be readily recognised. The family includes herbs with swollen nodes, as well as lianes and trees, often with oxidising wood. The leaves often dry dark grey and the venation is indistinct; the indumentum of the terminal bud is rich brown in color. The tubular, petaloid perianth with its induplicate-valvate or contorted aestivation is distinctive. The fruit is often ± drupaceous, or covered with sticky secretions and then ± exozoic; the fruit proper, an achene, is covered by the accrescent basal part of the perianth.

Molluginaceae [Halophytaceae + Cactaceae + Portulacaceae + Didiereaceae + Basellaceae + Hectorellaceae]: ?

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

• Molluginaceae are herbs or subshrubs with pseudoverticillate leaves that often have scarious "stipules", flowers that may be quite showy but which generally lack any corolline whorl, and seeds that lack an obviously long funicle.

Halophytaceae + Cactaceae + Portulacaceae + Didieraeceae + Basellaceae + Hectorellaceae: plants succulent; CAM +; secondary growth normal; phloem parenchyma cells with phytoferritin; Ca oxalate crystals in stem epidermis; P petaloid, pollen pantocolpate, similar fused columellae, etc.

Clade and taxon limits in this area are particularly difficult at present: Beware!

HALOPHYTACEAE A. Soriano - Vascular rays 0; A porose, endothecium with frame-shaped thickening on anticlinal walls, anther dehiscence by contraction of the connective, pollen cuboid; fruit a nutlet, several becoming enclosed by inflorescence axis. - 1/1. Argentina.

Cactaceae + Portulacaceae + Didiereaceae + Basellaceae + Hectorellaceae: 6-bp deletion in ndhf gene.

CACTACEAE Jussieu - Short shoots producing spines and hairs; stomata parallelocytic; P many, A many, 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.

Pereskia + Maihuenioideae + Opuntioideae + Cactoideae: delayed bark formation; stomata in stem epidermis.

2. Pereskia - 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 +.

Maihuenioideae + Opuntioideae: Leaves terete.

3. Maihuenioideae - 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 Schumann - Glochids with retrorse barbs; seeds surrounded by bony funicular "aril". - 5/225. S. Canada to Patagonia.

5. Cactoideae - Plant leafless; 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.

• Cactaceae can be recognised by their usually stout and very fleshy stems, axillary areoles of spines and hairs, and flowers that usually have an inferior ovary and many stamens and "petals"; the fruits are fleshy.

Portulacaceae + Didiereaceae + Basellaceae: median bracteoles [not subtending buds] present, enclosing flower; median P abaxial, no funicular obturator. Note: there is no evidence that this group forms a clade.

PORTULACACEAE Jussieu - 36/395 Talinum (40). ± World-wide, esp. W. America.

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

BASELLACEAE Rafinesque - G with one ovule. - 4/20. Pantropical, but mostly New World.

HECTORELLACEAE Philipson & Skipworth - 2/2. South Island of New Zealand and Kerguelen Island. NOTE: this is associated with some ex Portulacaceae in a clade that will perhaps be called Montiaceae Radlk.

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 J. Agardh - Stem parasites; sieve tube plastids lacking starch and protein inclusions; stem apex aboring; 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.

• Misodendraceae are stem parasites (hosts Nothofagaceae) that are recognizable by their stems, which are often thick, even when young, "giving the leafy species a rather elephantine appearance". The leaves may be reduced, and those on flowering shoots are different from those on vegetative shoots; the two kinds of shoots are sharply distinguished. The flowers are small, and the small, nutlike fruits are very distinctive with their three, long, plumose attachments.

SCHOEPFIACEAE Blume - pollen aperture development follows Garside's Rule [three pores at four points in the tetrad]. - 1/25. Tropical America, a few species in tropical South East Asia-Malesia.

LORANTHACEAE Jussieu - Collenchymatous zone below the embryo sacs; fruit with viscin; endosperm compound. - 68/950. World-wide.

• Loranthaceae are usually stem parasites that form secondary points of attachment along the stem. They are recognizable by their more or less opposite, often rather thick and brittle (especially when dry) leaves with entire margins and inconspicuous venation born on stems that are not jointed. Their 4-5-merous flowers are quite long, narrowly tubular and brightly colored - often red and/or orange - and lack much in the way of a calyx; the ovary is inferior and the fruits are fleshy. The flowers often have a bract/bracteoles at the base and immediately to one side of the ovary.

Opiliaceae + Santalaceae: P only.

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

• Opiliaceae may be recognised by their leaves, which are often 2-ranked, lack much in the way of venation, and have small bumps because of the cystoliths. The flowers are small, and are borne in inflorescences which are often almost catkin-like when young; there is only one perianth whorl. The ovary is superior.

SANTALACEAE R. Brown - 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.

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

SAXIFRAGALES Dumortier  

Ellagic acid +; cuticle waxes as clustered tubules [palmitone the main wax, at least in Paeoniaceae]; 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.

CERCIDIPHYLLACEAE Engler - Prophyll adaxial; stipule adaxial-petiolar; flower a pseudanthium, P 0, "A" and "G" several, G with abaxial suture, stigma long-decurrent; seeds winged. - 1/2. China and Japan.

• Cercidiphyllaceae are deciduous trees that may be recognised by the sharp distinction between the very long-lived short shoots and the long shoots; the leaves are more or less opposite, stipulate, and with palmate venation. The small flowers lack a perianth and the fruits are follicles.

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.

• Peridiscaceae are tropical trees with entire, stipulate leaves. The racemose inflorescences have small flowers, and the gynoecium has a central column and apical ovules; the style or styles are short. The capsular fruit has a single, large seed with a dark-colored but not very hard coat; the embryo is very small and embedded in cartilagineous endosperm.

DAPHNIPHYLLACEAE Müller Argoviensis - Iridoids +; leaves entire, stipules 0; P 0, G superior. - 1/10. East Asia to Malesia.

• Daphniphyllaceae are evergreen trees or shrubs that may be recognised by their rather weakly pseudoverticillate, exstipulate, entire leaves which are often glaucous beneath; the petioles of the leaves of a single innovation often vary considerably in length. The subcorymbose, racemose inflorescence with its small flowers with at most inconspicuous perianth is also distinctive.

HAMAMELIDACEAE R. Brown - Hairs stellate. - 27/82. Tropical to temperate, esp. East Asia to Australia, not South America.

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

2. Disanthoideae Harms - 1/1. E. China, Japan.

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

• Hamamelidaceae may be recognised by their often stellate or tufted hairs, their strongly stipulate leaves which often have ± palmate venation, their smallish flowers borne in rather dense inflorescences, and their bivalved, loculicidal and septicidal capsules with rather large seeds that have a hard, shiny, bicolored testa. The seed are sometimes forcibly ejected. The bases of the branches are often swollen.

ALTINGIACEAE Horaninow - Iridoids +; secretory canals with aromatic contents; stipules petiolar; inflorescence ± capitate; P 0, pollen polyporate. - 1/13. E. Mediterranean, East Asia to Malesia, Central America.

• Altingiaceae are monoecious, deciduous trees that may be recognised by their spirally-arranged, usually palmately-lobed leaves with petiolar stipules and by their capitate inflorescences. The buds have scales.

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.

• Paeoniaceae are perennial herbs or shrublets that are recognisable by their rather large, ± ternately compound, exstipulate leaves, and large flowers. The latter have sepals, petals, numerous centrifugal stamens, a prominent disc, and separate carpels. The fruit is a follicle and the seeds are fleshy.

Crassulaceae + Aphanopetalaceae + Tetracarpaeaceae + Haloragaceae + Penthoraceae + Saxifragaceae + Iteaceae + Pterostemonaceae + Grossulariaceae: 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; testa sinuate-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.

• Crassulaceae are recognisable by their succulent herbaceous or soft-stemmed habit, and their flowers, which have the same number of sepals, petals and carpels; the latter are more or less free and have nectariferous scales at their bases. The fruits are usually follicles.

Aphanopetalaceae + Tetracarpaeaceae + Penthoraceae + Haloragaceae: ?

APHANOPETALACEAE Doweld - C 0, one apical ovule/carpel; fruit a nut. - 1/2. W. and E. Australia.

• Aphanopetalaceae are scrambling shurbs with prominently lenticellate stems, opposite and more or less serrate leaves with minute "stipules", and axillary inflorescences. the flowers are apetalous and have a spreading calyx that enlarges in fruit; the fruit itself has a single seed.

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

TETRACARPAEACEAE Nakai - Fibrous endothecium 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.

• Penthoraceae are rather fleshy herbs with spiral, entire leaves. They may be recognised by their few-branched, monochasial cymose inflorescence, and often apetalous flowers with partly connate carpels with submarginal styles. The fruits are distinctive since the free part of each carpel splits circumscissilely at the base.

HALORAGACEAE R. Brown - 8/145. World-wide, especially Australia.

• Haloragaceae are rather small, usually monoecious, often more or less herbaceous plants. Their leaves are serrate or deeply lobed, small, and often opposite or whorled, and their small flowers (Haloragodendron is an exception) have penicillate stigmas.

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

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

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

• Iteaceae may be recognised by their stems with chambered pith, spirally-arranged serrate leaves with superposed axillary buds, paniculate to racemose inflorescences with rather small flowers, and septicidal fruits the valves of which often remain attached by the stigma. The flowers have a hypanthium, the corolla is valvate, and there are only two carpels.

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

• Pterostemonaceae look rather like Rosaceae, both being woody and having spiral, serrate, stipulate leaves, although those of Pterostemonaceae tend to be notably shiny. However, the distinctive conical to peltate glandular hairs, the exudate of which produces the shiny leaf surface, and the flowers, each with a whorl of five stamens that have winged and toothed filaments and a whorl of five staminodes, allow the family to be recognised.

Grossulariaceae + Saxifragaceae: G [2-3].

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

• Grossulariaceae may be recognised by their lobed, spiral leaves with broad bases, and flowers with a well-developed hypanthium, relatively large and colored sepals and smaller petals, a usually bicarpellate gynoecium, and parietal placentation; the fruit is baccate.

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

• Saxifragaceae can be recognised by their herbaceous habit and gland-toothed leaves; stipules, if present, are often on the petiole near the base. The flowers often have 2-3 carpels that are basally connate, a disc is common, and the fruit is a follicle or septicidal capsule.

CYNOMORIACEAE (Schott & Endlicher) Lindley - Echlorophyllous root parasite; 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 + [multicellular, spherical, with an apical stoma]; leaves with glandular teeth; 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

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

• Vitaceae may be recognised by their caducously stipulate leaves with palmately compound, -lobed or -veined blades and often rather coarse teeth, cymose inflorescences, and smallish flowers with valvate petals. The stamens are equal in number to and opposite the petals, the usually 4-ovulate, bicarpellate gynoecium has more or less sessile stigmas, and the berry often has ruminate seeds. The stems are often strongly lenticellate, with the bark ultimately flaking, and the leaf blades are very brittle when dry, the fine details of the venation being obscure. Leaf-opposed tendrils or inflorescences are common in the family.

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

Unplaced:

PICRAMIACEAE Fernando & Quinn - C₁₈ acetylenic acids [tariric, petroselenic], anthraquinones, anthracenone moieties linked to C₅-sugar derivatives +; 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. USA, 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.

• Picramniaceae are often rather small trees with bitter or very bitter-tasting bark and spirally-arranged, compound leaves; the leaflets alternate along the rachis and the bark is bitter. The small flowers are born in a raceme, and the stamens are equal to and opposite the petals.

CROSSOSOMATALES Reveal  

?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. - 3(?)/45. North temperate, tropics in America (to Bolivia) and Malesia.

• Staphyleaceae are shrubs to trees that may be recognised by their opposite, odd-pinnately compound leaves with serrate leaflets and little "glands" at the articulations (the latter often collapse when dry), the stipules are cauline and are sometimes interpetiolar. The flowers have rather petaloid sepals and usually two to three carpels.

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.

• Guamatelaceae are sprawling shrubs with opposite, stipulate, serrate leaves that have palmate venation and a densely whitish tomentose lower surface. The flowers have sepals and petals almost equal in size, ten stamens, and three carpels connate only by the tips of their styles. The fruit are follicles and are almost enclosed by the persistent sepals.

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

STACHYURACEAE J. Agardh - Stomata paracytic; inflorescence long-racemose; hypanthium 0, nectary at base of G. 1/5. South East Asia.

• Stachyuraceae are small trees or shrubs with closely serrate, stipulate leaves and rather long, pendulous, racemose inflorescences. The sepals and petals of the four-merous flowers are similar, and the fruit is a berry.

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

• Crossosomataceae are rather small-leaved, much-branched deciduous shrubs that are otherwise vegetatively nondescript. The flowers have separate sepals, petals and carpels and a more or less well developed hypanthium. The follicles open down their abaxial surfaces as well and the seeds have a fimbriate aril.

Aphloiaceae + Geissolomataceae + Ixerbaceae + 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.

• Aphloiaceae are undistinguished evergreen shrubs or small trees with distichous, serrate leaves and often persistent stipules. The axillary flowers have a hypanthium, an undifferentiated perianth, many stamens, and a single carpel. The fruit is a berry. Herbarium specimens have a yellowish, crusty material associated with them, rather like some Thymelaeaceae.

Geissolomataceae + Ixerbaceae + 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.

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.

Ixerbaceae + Strasburgeriaceae: 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.

IXERBACEAE Grisebach - Stipules 0; A = and opposite K, pollen 4-5-colporate; carpels also splitting adaxially, valves woody, reflexed. - 1/1. New Zealand.

• Ixerbaceae are evergreen trees with pseudoverticillate and strongly toothed but estipulate leaves. The inflorescences are terminal and the flowers are rather large and have five spreading sepals and petals, ten stamens with thick connectives, and a conspicuous ovary gradually tapering into the style and with a spreading, lobed nectary at its base. The pointed capsular fruit has a few shiny black seeds with a conspicuous red aril.

STRASBURGERIACEAE Solereder - Stipules intrapetiolar, connate basally; flowers axillary; hypanthium 0, nectary on G, one ovule/carpel; fruit rather dry, indehiscent; n = ca 250. - 1/1. New Caledonia.

• Strasburgeriaceae are evergreen trees that may be recognised by their rather large, spiral and toothed leaves that have intrapetiolar stipules. The flowers are large and axillary and have spiral sepals, five petals and ten stamens with long anthers attached their lengths to the thick connective. The stoutly-pointed fruit is indehiscent.

GERANIALES Dumortier

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. - 6/805. Temperate and warm temperate, esp. southern Africa.

• Geraniaceae are often herbs with jointed stems; the stipulate leaves have palmate venation or are palmately or pinnately compound. The inflorescence is cymose, and all parts of the flower apart from the carpels are free. The sepals are aristate, the androecium is obdiplostemonous, the carpels are usually beaked, there are few (1-2(-12)) ovules per carpel, and the embryo is curved.

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, micropyle endostomal, funicle with hairs, 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.

• Myrtales may be recognised by their often flaky bark; opposite leaves with colleters, at most small stipules and strong intramarginal veins; flowers with petals that are clawed or at least very narrow at the base, a more or less inferior ovary, and a usually nectariferous hypanthium. The sepals are often valvate and the stamens are usually incurved in bud.

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.

• Combretaceae may not be very easy to recognise, although like other Myrtales they often have opposite leaves and the bark is often flaking. Trees quite often look rather pagoda-like, the branching following the Terminalia pattern, the branches consisting of flattened sprays of leaf rosettes. The leaves are entire and may have pellucid dots, surface glands (also on the petiole), or domatia. The indumentum is commonly adpressed and distinctive enough under the microscope; nodes are unilacunar. The flowers are often rather small and massed in spicate inflorescences, the hypanthium can be quite long, but the corolla is small or absent. The ovary is inferior, and the single-seeded fruit is often flattened and/or winged.

Onagraceae + Lythraceae: tannins often not abundant; 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; 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; fruit a capsule opening down the sides. - 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.

• Onagraceae are herbs to shrubs that are recognisable by their usually opposite leaves with small stipules and the often 4-merous flowers with an inferior ovary and a well-developed hypanthium. The calyx is rather thin and reflexed, there are up to eight stamens, and the pollen has viscin threads, so it is conspicuously clumped.

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

• Lythraceae are herbs to trees that are recognisable by their flaky bark, opposite leaves that lack gland dots, flowers with a ribbed hypanthium or K + C tube, a rather thick, valvate calyx, clawed petals that are crumpled in bud, and often 10 or more stamens of unequal lengths. The ovary is more or less superior, and the fruit is usually a capsule.

Rest: ?

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

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

• Vochysiaceae are recognisable by the combination of rather ordinary Myrtalean vegetative body (although some taxa have large, flat, stipular glands) combined with the distinctive monosymmetric or asymmetric flowers with a spurred, sometimes cochleate K and a single stamen.

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 - 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.

• Myrtaceae are recognisable by their flaky bark, their entire, punctate leaves, and their imbricate calyx, often rather fugaceous corolla, and androecium with numerous stamens each often with an apical gland. In fact, in leaf, flower and fruit there is considerable variation. Many taxa have opposite leaves with a well-developed intramarginal vein; there is a group of genera with spiral leaves that dry a distinctive, dark, almost purplish color. Many genera have inferior ovaries and baccate fruits, while another large group of genera has ± superior ovaries and capsules.

Melastomataceae + Crypteroniaceae + Alzataeaceae + Penaeaceae + Oliniaceae + Rhynchocalycaceae: (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, pollen with pseudocolpi; radicle bent. - 188/500. Pantropical (warm temperate).

?name of clade - Lamina with veins diverging at the base, tertiaries scalariform; A held on one side of the flower. - 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.

• Melastomataceae are recognisable by their opposite, entire, exstipulate leaves with three or more strong veins ascending from at or near the base; tertiary veins are conspicuous, and run at right angles to the midrib. Hair types are notably diverse. The flowers with their contorted and often somewhat stridently pink petals and porose anthers which invert during development and have an elaborate, expanded connective, are also distinctive. In old fruits, the walls often distintegrate, leaving vertical vascular bundles behind.

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

• Memecyloideae have opposite, entire leaves and swollen nodes; the venation is variable, but indumentum is absent or consists of uniseriate hairs. The blade can be minutely punctate when held up to the light, or the surface may be minutely lineate - this is because of the sclereids common in the family (the surface may also dry minutely bumpy). The rather small flowers have inferior ovaries, but the parts are otherwise free, the calyx is valvate and there are twice as many stamens as petals; the anthers dehisce by slits and there are glands on the connectives. Most species are plants of the understorey.

Crypteroniaceae + Alzataeaceae + Penaeaceae + Oliniaceae + Rhynchocalycaceae: stems square, nodes swollen, stipules minute; A = and opposite C, endothecium ephemeral; fruit a capsule; exotestal cells periclinally elongated, endotegmen not fibrous.

CRYPTERONIACEAE A.-L. de Candolle - 3/10. South East Asia, Malesia, Sri Lanka.

Alzateaceae + Rhynchocalycaceae + Penaeaceae + Oliniaceae: style stout.

ALZATEACEAE S. Graham - C 0, pollen pseudocolpi 0 [faint], placentation parietal and with an incomplete septum, embryo sac bisporic, Allium-type. - 1/?2. Costa Rica to Peru.

Rhynchocalycaceae + Penaeaceae + Oliniaceae: leaves with glandular tips; n = 10.

RHYNCHOCALYCACEAE L. A. S. Johnson & B. G. Briggs - Anthers with 4 loculi opening separately. - 1/1. South Africa

Penaeaceae + Oliniaceae: plants not Al accumulators; non-hydrolysable tannins +; hypanthium well developed, pollen grains psilate, foot layer and tectum thick.

PENAEACEAE Guillemin - 7/20. South Africa, the Cape.

OLINIACEAE Arnott - K ± spatulate, C concave, thick, hairy, pollen heteropolar; fruit drupaceous. - 1/5. Africa, St. Helena.

EUROSID I: endosperm scanty.

Unplaced:

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.

• Huaceae are evergreens, usually trees, that are recognizable by their 2-ranked leaves the blades of which have a very fine, boxy reticulum; there are small glands at the very base of the lamina. The cauline stipules are early deciduous. The plant smells of garlic.

ZYGOPHYLLALES Chalk

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.

• Krameriaceae may be recognised by their small, spiral, often elliptic, entire leaves and their distinctive, rather pea-like flowers. These are inverted, with the odd petal adaxial, and have four or five large elliptic sepals that are petaloid internally; the three strongly clawed adaxial petals are more or less connate by their bases and the usually four fertile stamens are also adaxial. The one-seeded fruits have retrorsely-barbed spines.

ZYGOPHYLLACEAE R. Brown - 22/285. Dry and warm temperate, also tropical.

• Zygophyllaceae are a variable group growing in more or less arid and saline conditions throughoutt the world. They are herbs to trees with compound usually stipulate and opposite leaves; the nodes tend to be swollen. The flowers have all parts free apart from the carpels; there are usually ten stamens that often have scales at the base. Some aspects of floral morphology, e.g. obdiplostemony, the stigma, are reminiscent of Geraniales.

CELASTRALES + MALPIGHIALES + OXALIDALES: seed exotegmic.

CELASTRALES Baskerville

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; A 10, of two lengths, ± connate basally, anthers basifixed, nectary on inside of staminal tube, G [2-3], 2 collateral apical epitropous ovules/carpel; fruit a septicidal capsule, endocarp distinct, columella persisting. - 2/2-3. E. Africa, Central and South America.

• Lepidobotryaceae may be recognised by their entire, simple but articulated leaves with both stipules and stipels. The inflorescence often seems to be opposite the leaves because it is evicted by the growth of a robust axillary shoot. The flowers are small, with free sepals and petals and ten basally-connate stamens; the antepetalous stamens have longer filaments than the antesepalous whorl. The mesocarp, with its distinctive ladder-like fibers arranged radially, separates from the horny endocarp.

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.

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.

• Parnassiaceae are herbs that may be recognised by their serrate leaves that often have reddish lines or dots when dry. The flowers are radially symmetrical, petals, when present, are free, there are only five stamens, and the ovary has parietal placentation. There are five staminodes, those of Parnassia being large and particularly distinctive.

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

• The family is vegetatively variable - Microtropis may have oppposite, entire, exstipulate leaves, while spiral and opposite leaves on the one plant are not infrequent, as in Catha and Salacia. However, opposite, serrate leaves are common and the stipules are small and quite often fringed; strongly ascending veins are prominent on dried specimens. Thin, spiral threads are often evident when the lamina is torn transversely. The inflorescences are often cymose, and the flowers, with their usually very broad disc in which the ovary is more or less immersed and five or fewer stamens, are often distinctive. The fruit varies greatly, but it is frequently a loculicidal capsule with winged or arillate seeds. The colleters may turn black, as in Paxistima.

MALPIGHIALES Martius

Leaf margin toothed [teeth with a single vein running into a congested ± deciduous apex]; stigma dry; exotegmen fibrous. - 38 families, 716 genera, 15935 species.

Achariaceae + Violaceae + Salicaceae + Lacistemataceae + Goupiaceae + Malesherbiaceae + Turneraceae + Passifloraceae: 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; seed coat thick, testa vascularised. - 30/145. Pantropical.

• Achariaceae can be recognised by their stipulate leaves, often entire leaf blades and pulvinate petioles, and flowers in which the number of sepals and petals is often clearly not the same, and/or the perianth is not simply biseriate, and/or the petals have basal, adaxial scales. There is no nectary, and the anthers are usually long. The fruits are often quite large, the placentation is parietal, and the seeds often have conspicuous vascular bundles in the coat.

VIOLACEAE Batsch - Abaxial C spurred, anther connective well-developed. - 20/800. World-wide; woody taxa esp. in the lowland tropics.

• Violaceae may be recognised by their serrate, stipulate leaves which often dry yellowish and with raised reticulum, their flowers, which are often spurred and have stamens with very well-developed connectives, and their fruit, which is often a distinctive loculicidal capsule, the rather large seeds being dispersed explosively as the fruit wall closes round them and squeezes them out. They are herbs, trees or lianes.

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

• Salicaceae can be recognised by their alternate, serrate, stipulate leaves and flowers with many often spreading stamens and a commonly three-carpellate gynoecium with parietal placentation and either separate styles or a single, long, persistent style; the fruit is often a capsule. When there are both sepals and petals, the two whorls are in a regular alternating arrangement, and the nectariferous disc or nectar glands and often outside the stamens. A number of taxa have pellucid foliar glands.

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

• Lacistemataceae are woody plants that have stipulate and usually serrate leaves. The small flowers are borne in rather dense racemes to spikes and have an inconspicuous perianth and a single anther whose two thecae are often well separated and even stipitate. The dehiscent fruit is small, with one, sometimes to three, apical seeds.

Turneraceae + Malesherbiaceae + Passifloraceae [= Passifloraceae s.l.]: 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); funicular aril +, endotestal cells large, exotegmen palisade, endotegmen persistent.

Turneraceae + Malesherbiaceae: leaves spiral; micropyle bistomal, style branched; x = 7.

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

• Turneraceae are herbs to woody plants that may be recognised by their often very hairy, serrate, exstipulate leaves with strong secondary venation. The flowers, with their calyx and corolla together forming a tube, contorted and deliquescent yellow or sometimes red corolla, parietal placentation, and often bifid, fringed stigmas, are distinctive. The loculicidally capsular fruits have arillate seeds with a minutely and often regularly reticulate testa.

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

• Malesherbiaceae are are more or less herbaceous, foetid and often densely glandular-hairy plants with distinctive flowers. The calyx and corolla together form a long tube; the sepals and petals are both valvate. There is an androgynophore, the floral tube is persistent in fruit, and the styles are often subapical on the capsule.

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

• Many Passifloraceae are tendrillar vines or lianes; the leaves are alternate, stipulate, often with palmate venation, and often with glands on the lamina or petiole. The tendril is often oblique to the axis; a flower may be lateral to it, or paired flowers are borne on it or at its base. The axillary buds are superposed, and the lower bud produces the inflorescence/tendril. Floral features are, however, more important for the recognition of the family. The calyx and corolla are often similar and fused to each other at the base, and there is usually a complex corona. There is a gynophore or androgynophore, five large and versatile anthers, and a three-carpellate gynoecium with parietal placentation. The fruit is a berry, and the often flattened seeds are arillate.

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.

• Goupiaceae may be recognised by their two-ranked stipulate leaves with ascending secondary veins and scalariform tertiaries. The inflorescence is umbellate and the petals are relatively long and inflexed at the apex. The gynoecium has basal-axile placentation and separate styles; the fruit is a single-seeded drupe.

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

LOPHOPYXIDACEAE (Engler) H. Pfeiffer - Liane with leaf tendrils; secondary thickening with included phloem; fruit a 5-winged samara. - 1/1. Malesia to the Solomon and Caroline Islands.

• Lophopyxidaceae are tendrillate lianas with quite strongly serrate leaves that have pinnate venation; the flowers are small and are borne in tight clusters along the branched inflorescences. The one-seeded fruit has five wings.

PUTRANJIVACEAE Endlicher - Glucosinolates +; plant dioecious; 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.

• Putranjivaceae are trees to shrubs that may be recognised by their 2-ranked often rather coriaceous leaves that are asymmetrical at the base and dry a greyish color; they frequently taste peppery or like an inferior radish when fresh, although the taste may take a little time to develop (mustard oils!). The fasciculate flowers are frequently rather small and the single-seeded drupe is often crowned by the persistent, flap-like stigmas.

Erythroxylaceae + Rhizophoraceae: tropane [hygroline] and pyrrolidine alkaloids +; sieve tube plastids with protein crystalloids; mucilage cells +; stomata paracytic; leaves opposite, involute, colleters +, stipules enclosing the terminal bud; inflorescence cymose; 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.

• Erythroxylaceae are shrubs to trees that can be recognised by their entire leaves, stipules that are at least in part interpetiolar, and usually fasciculate inflorescence with rather small flowers; both calyx and filaments persist at the base of the fruit. Erythroxylum itself has intrapetiolar stipules, there are often longitudinal markings on the leaves, the filaments are often of two different lengths and are basally connate, and the fruit is a drupe.

RHIZOPHORACEAE Persoon - 16/149. Pantropical.

• Rhizophoraceae are shrubs to trees that may be recognised by their opposite often serrate leaves and large interpetiolar stipules. The flowers are quite often other than 5-merous, the calyx is thick and valvate, the petals are often lobed or finged, each enclosing small groups of stamens, and the ovary is ± inferior. Gynotrocheae and Rhizophoreae have aerial prop roots.

LINACEAE Perleb - 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.

IXONANTHACEAE Miquel - Stomata paracytic; pollen with supratectal spines, ovules apical. - 4-5/21. Pantropical.

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

• Ctenolophonaceae may be recognised by their opposite, entire leaves with interpetiolar stipules and their terminal inflorescences with rather elongated flower buds. The rounded sepals are persistent and accrescent, and the grey-drying and closely-ribbed fruit is distinctive.

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

• Although hardly very distinctive, Centroplacaceae are evergreen trees with serrate and stipulate leaves that can be recognised by their branched inflorescences of small flowers; the staminate flowers have only five stamens with a well-developed connective and two oblique-apical slits.

HUMIRIACEAE A.-L. de Jussieu - 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.

• Humiriaceae are evergreen trees that may be recognised by their blackish-drying, short-petiolate, toothed leaves which elongate while still rolled up and often have longitudinal lines down the blade; the buds on growing shoots are long-pointed. The young twigs are angled. The cymose inflorescences have rather small flowers, and the anthers have conspicuously prolonged connectives and the sepals are broadly rounded. The fruit is a drupe and has a ridged, operculate stone that often has cavities containing almost resin-like material.

PANDACEAE> Engler & Gilg - 3/15. Tropics, Africa to New Guinea.

• Pandaceae are evergreen trees with stipulate leaves and often very strongly plagiotropic branches that have even been confused with compound leaves. The flowers are small and inconspicuous, the plants are dioecious, and the fruit is a drupe.

IRVINGIACEAE (Engler) 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.

• In Irvingiaceae, the leaves, with their longitudinal markings, their large, deciduous, intrapetiolar stipules that enclose the prominent, pointed terminal bud, and their rather closely parallel secondary venation, are distinctive. The axillary buds are also prominent, and prophylls are basal on the shoots. The pedicels are articulated at the junction with the axis; the thin calyx persists on the fruit and is conspicuously reflexed, and the disc is very obvious. The drupes often have radial fibers.

Clusiaceae + Bonnetiaceae + 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 Lindley - Exudate +. - 27/1050. Pan tropical.

1. Kielmeyeroideae Engler - 14/465. Throughout the tropics.

2. Clusioideae Engler - Hypocotyl massive, cotyledons minute. - 13/590. Throughout the tropics.

• Clusiaceae may be recognised by their usually opposite, exstipulate leaves with entire margins and usually quite obvious glands or canals; there is often some exudate from the cut twig. The flowers have free sepals and petals and numerous stamens, quite often in fascicles; the anthers may bear various kinds of glands. The fruit, when dehiscent, opens along the septal radius.

• Kielmeyeroideae are usually trees. Their leaves have pellucid dots, rarely canals, and the androecium is not obviously fasciculate, the anthers sometimes having large, apical glands. The seeds range from small to large; when large, they are made up almost entirely of two huge cotyledons.

• Clusioideae range from shrubs or trees, some are epiphytic; many are dioecious. The leaves usually have canals and the androecium is sometimes obviously fasciculate. The seeds are small to large, arils being known from many New World taxa, and the embryos are often apparently undifferentiated - the cotyledons are small to almost absent and the hypocotyl is enormous.

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

• Bonnetiaceae may be recognised by their long-pointed buds, usually rather closely-set, spiral, short-petiolate and minutely-serrate leaves. The moderate-sized flowers are borne in cymose inflorescences, the corolla is contorted, the stamens are numerous, and the septicidal capsules contain many small seeds.

Hypericaceae + Podostemaceae: canals, cavities or individual cells with exudate; ovules tenuinucellate, stigma papillate.

HYPERICACEAE Jussieu - 9/560. World-wide.

• Hypericaceae include numerous herbs or shrubs. Their leaves are opposite and with pellucid or dark dots; the androecium is often fasciculate; the stigma is sometimes obviously papillate; and the seeds are small (usually less than 4 mm long).

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; 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; capsule ribbed; hypocotyl 0. - 43/260. Pantropical.

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

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

• Podostemaceae are the quintessential aquatic angiosperm with a vegetative body so highly modified that conventional terms cannot be used to describe it - the plant body of many taxa is best described as thalloid or fucoid, although some species look like a demented apiaceous plant with 2-ranked leaves. Many species are annual, and flowers or groups of flowers are often enveloped in a spathella; the capsule is septicidal, and the small seeds are often mucilaginous.

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.

• Malpighiaceae are a distinctive family, with their usually opposite, entire leaves with paired stipules (these vary in position). The unicellular hairs are T-shaped, often a characteristic brown, and there are often conspicuous flat glands on the leaf blade or petiole. The flowers are characteristic: 4 or 5 sepals often have paired glands, the petals are strongly clawed and are crumpled in bud, and there are three carpels, each with a single ovule. The sepals and filaments often persist at the base of the fruit, which is often a samara with a variable number of wings.

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

• Elatinaceae are often herbaceous plants of moist habitats with opposite, more or less toothed leaves with small, paired, scarious stipules. The cymose inflorescences have small flowers with papillate stigmas.

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 + Medusagynaceae: ?

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; testa with vascular bundles, lacking layer of small crystalliferous cells, fibrous exotegmen 0; endosperm 0. - 9/290. Tropical, especially Brazil.

3. Sauvagesioideae Beilschmied - 16/82. Pantropical, only 2 spp. in Africa, most genera South America.

• Ochnaceae are recognisable by their alternate, often coriaceous leaves with rather sharply serrate margins, more or less brochidodromous venation - the secondary veins can be very closely parallel - and rather scaley stipules that have strong, parallel veins and also may have fringed margins. The rather scarious calyx is also distinctive, the anthers are often porose and the style sometimes gynobasic.

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.

• Medusagynaceae are evergreen trees that have a distinctive fibrous bark like that of Juniperus. The leaves are opposite, toothed, and with very strongly reticulate venation. The flowers, with their contorted petals, numerous stamens and carpels with marginal stylodia are distinctive, as are the fruit, the valves of which pull away from the central columella, except at their apices.

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

• Quiinaceae are evergreen trees that may be readily recognised, even when sterile, by their opposite or whorled, sometimes compound (especially in seedlings), strongly stipulate leaves with toothed margins; the secondary veins are well-developed, the fine venation is like the strokes of a paint brush, and the stipules are more or less persistent. There are well-developed mucilaginous canals. Both stipules and sepals are pubescent and the often baccate fruits are striate when dry.

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, polythecate, pores terminal, microsporogenesis successive; carpellate flowers: G inferior, placentation laminar-parietal, ovules tenuinucellate, nucellar cap +, nucellar epidermis persists, stigma on outer margin or underside of disc; seed in two parts, that covered by the testa not enveloping the embryo, exotegmic cells with U thickenings; endosperm slight, embryo undifferentiated. - 3/20. Central and South America, the Antilles, Near East, W. Malesia, Australia.

• Rafflesiaceae are stem or root parasites lacking both rhizomes and roots. The flowers are large to massive, imperfect and with a uniseriate, connate perianth; there are 12-40 extrorse stamens adnate to a central column in the staminate flowers and the carpellate flowers have an inferior ovary.

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.

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.

• Euphorbiaceae may be recognised by the leaves on the one branch often being variable in size and/or shape; the secondary veins are often ± palmate, the margins toothed, and there are often glands. Cauline stipules are common, as is latex (the latter not in Acalyphoideae). The plants are mono- or dioecious, the flowers are small, the perianth usually inconspicuous, there are usually three carpels with prominent stigmas, and the fruits have a distinctive and persistent columella, large and often carunculate seeds (one per loculus), and explosive dehiscence.

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.

• Phyllanthaceae have a variety of growth forms, but can be recognised by their often finely-cracking bark, absence of latex, often 2-ranked, stipulate and pinnately-veined leaves lacking glands, and explosively dehiscent fruit that have a persistent columella; there are two ovules in each carpel.

PICRODENDRACEAE Small - 27/85. Many small genera, tropical, esp. New Guinea - Australia - New Caledonia (Caletieae) or America plus Africa - Madagascar (the rest).

• Picrodendraceae are vegetatively rather heterogeneous, although they often have opposite, toothed and stipulate leaves with pinnate venation and deciduous glands capping their teeth. The capsules are like those of Euphorbiaceae and the seeds may be carunculate.

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, 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.

• Balanopaceae look like a Myrica, but they lack its distinctive glandular hairs. The leaves are toothed, albeit sometimes minutely, and are very coriaceous. The staminate flowers are in catkins, the carpellate flowers are single and surrounded by a cupule of spirally-arranged bracts; the flowers have no more than a rudimentary perianth. When dry, the cupulate, drupaceous fruits look somewhat like acorns; they have 2-3 stones.

Trigoniaceae + Dichapetalaceae + Chrysobalanaceae + Euphroniaceae: vestured pits +; mucilage cells +; stomata paracytic; leaf margins entire (surface glands or glandular hairs +); (flowers obliquely monosymmetric); ovules tenuinucellate, 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. - 5/28. Central and South America, Madagascar, W. Malesia.

• Trigoniaceae may be recognised by their simple, entire, stipulate leaves that have whitish indumentum below and by their obliquely monosymmetric flowers that are papilionaceous in general appearance.

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

• Dichapetalaceae are trees to lianes that may be recognised by their lenticillate twigs and stipulate, often dark greyish-drying leaf blades with brochidodromous venation; this last is prominent on the abaxial surface, where there are also often flat glands. The inflorescences are cymose and ± obviously epiphyllous; the small flowers have deeply bifid petals which dry blackish. The fruit is a drupe that is often flattened and/or lobed and is covered by short, dense, erect hairs which often have a shiny, almost golden appearance; it is rarely dehiscent.

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.

• The family can be recognised by its strongly lenticillate twigs and 2-ranked, stipulate, and nearly always entire leaves; these either dry blackish grey and/or have well-developed scalariform tertiary venation. There are often broad, flat glands on the abaxial surface of the lamina, and the petiole is often short and more or less swollen at one or both ends. The flowers have a hypanthium, there are often many long stamens, and the style is more or less gynobasic, the ovary being borne on one side of the hypanthium. The one-seeded fruit is usually relatively large and the endocarp is often hairy inside.

EUPHRONIACEAE Marcano-Berti - 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. - 1/1-2. The Guyana Shield, South America.

• Euphroniaceae are woody plants with entire, spiral, stipulate leaves the lower surfaces of which are covered with whitish indumentum; there are marginal abaxial glands near the base. The monosymmetric flowers are distinctive in having a short hypanthium and only three free petals; the fruit is a columellate, septicidal capsule.

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.

• Caryocaraceae are evergreen trees to shrubs that may be recognised by their trifoliolate, stipulate and usually serrate leaves, terminal racemose inflorescences, and large flowers with very long, radiating stamens. The fruits are drupes and the seeds are reniform.

OXALIDALES Heinze

Vessel element type?; mucilage cells +; stomata ?; 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; leaves odd-pinnate, 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.

• Connaraceae are mostly woody lianes that twine in the apical parts of the innovations. They may be recognised by their pinnately compound, exstipulate leaves with transversely-ridged petioles, and ± ridged or keeled follicles with black and contrasting orange to red seeds. The follicles are usually single and swell to almost mature size well before the seeds are developed; the seeds are partly squeezed out of the follicle. The small, radial flowers with stamens of two lengths are rather undistinguished.

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.

• Oxalidaceae may be recognised by their usually compound leaves with pulvinate leaflets; their leaflets are entire and often have subpalmate venation. The inflorescence is either cymose or has cymose branches, the flowers are heterostylous, the corolla is clawed, and soon withers or is deliquescent, the stamens are obviously of two lengths, and the styles have capitate stigmas. The fruit is often a more or less ribbed loculicidal capsule.

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

CUNONIACEAE R. Brown - Leaves opposite, odd-pinnate, 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.

• Cunoniaceae may be recognised by their opposite, usually odd-pinnately compound leaves with serrate leaflets and well-developed often intrapetiolar stipules. The flowers are often quite small and the filaments are notably longer than the petals. The fruit, when a capsule, is septicidal.

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, odd-pinnate, 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.

• Brunelliaceae may be recognised by their dense brown indumentum, their robust, rather flattened twigs, their opposite, usually odd-pinnately compound leaves with strongly serrate stipellate leaflets, and their rather small, paired cauline stipules (cf. many Cunoniaceae!). The flowers are apetalous and have separate carpels and long-decurrent stigmas, and the fruits are follicles containing one or two shiny seeds that have an aril-like raphe down one side.

CEPHALOTACEAE Dumortier - Insectivorous herbs; leaves ascidiate or not, 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.

• Cephalotacaceae are insectivorous herbs that may be recognised by their rosette of leaves, some of which are fluted pitchers with lids (the leaves are reported to look like "vegetable hedgehogs" when young!), and scapose inflorescences with cymose branches. The six-merous flowers are small, lack petals, and have free carpels.

ELAEOCARPACEAE Candolle - 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.

• Elaeocarpaceae are woody plants that can be recognised by their leaves that tend to turn red or orange red, sometimes yellow, as they die; the blades are often serrate and the petiole can be swollen at one or both ends. Terminalia branching is common. The moderate-sized, pendulous flowers are borne in often racemose inflorescences. Both calyx and corolla are valvate (the latter not always), the petals, when present, are often fringed, there is often a very prominent disc that is usually outside the stamens, the basifixed anthers often have rather short filaments and dehisce by pores or short slits (these may elongate), and the fruit is either a drupe or berry or an often muricate capsule with arillate and/or hairy seeds.

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

FABALES Bromhead

Ellagic acid 0; embryo green. - 4 families, 754 genera, 20055 species.

POLYGALACEAE Hoffmannsegg & Link - Median abaxial C clawed, ± boat-shaped/keeled, A 8, median adaxial A 0, pollen polycolporate; K caducous; 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

• Polygalaceae are woody to herbaceous plants that may be recognised by their often yellowish-drying simple and entire leaves; there are quite frequently paired glands at the nodes or glands or domatia on the leaf blades. The flowers are usually monosymmetric and are more or less papilionaceous; they may have either five rather large sepals, or two of the sepals may be much enlarged and coloured (the "wings" of the papilionaceous flower). There are two or more carpels.

Surianaceae + Quillajaceae + Fabaceae: carpels free.

SURIANACEAE Arnott - 5/8 Mostly Australian, also Mexico, Suriana pantropical.

Quillajaceae + Fabaceae: hypanthium +, C clawed, many ovules/carpel; fruit a follicle, also opening abaxially; tegmen crushed; cotyledon investing radicle.

QUILLAJACEAE D. Don - Nodes 1:3; 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; fruit strongly lobed; seeds winged. - 1/3. Temperate South America.

• Quillajaceae are vegetatively rather undistinguished with their spiral, serrate, stipulate leaves, although the conspicuous teeth appear to be hydathodal. They may be readily recognised by their remarkable flowers in which the antesepalous stamens are borne on the outer edge of the disc some way up the sepals and the antepetalous stamens are near the base of the ovary. The petals are clawed and the strongly lobed fruits open down both the inner and outer surfaces of the lobes and contain winged seeds.

FABACEAE Lindley//LEGUMINOSAE Jussieu - Non-protein amino acids and lectins [hemagglutinins] esp. in seeds +; G 1, many serial ovules/carpel; fruit a follicle, also dehiscing abaxially; exotesta palisade, linea lucida separating much thickened outer anticlinal walls from the rest, mesotesta of stellate cells; endosperm ± starchy [Schleimendosperm]; rpl22 gene absent. - 730/19400. World-wide.

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

"Caesalpinioideae" + Mimosoideae + Faboideae: leaves bipinnate; vestured pits +; hour-glass cells [below palisade exotesta] +.

2. "Caesalpinioideae" - 160/1930. Predominantly tropical.

3. Mimosoideae - Sieve tube plastids also with fibres; extrafloral nectaries common; flowers of inflorescence develop together; 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.

4. Faboideae - Isoflavonoids [pterocarpans and isoflavans]+; sieve tubes with spindle-shaped non-dispersive protein bodies; adaxial C external; embryo with well-developed suspensor [?basal genera], bent, cotyledon not investing radicle, cotyledon areoles common. - 476/13855. Esp. (warm) temperate, but world-wide.

• Fabaceae may be recognised by their compound, stipulate leaves whose leaflets are entire and often have wrinkled pulvini, and flowers with a single carpel, free petals (or two may be connate), and stamens either all similar in length, or if dissimilar, not regularly alternating longer and shorter.

• "Caesalpinioideae" and Cercis and its relatives are woody plants that may be recognised by their usually once-compound leaves and monosymmetric flowers with the odd petal adaxial and inside the others in bud; the hypanthium is often well-developed and the seeds have a pleurogram.

• Mimosoideae are also mostly woody plants that have leaves that are often bicompound and with extrafloral nectaries. The flowers are rather small and are borne in groups, all flowers opening more or less simultaneously; the long, colored filaments form the visually attractive part of the inflorescence. Both the calyx and corolla are valvate, and both (and often the bases of the filaments) are connate. The seeds have both a pleurogram and a linea fissura.

• Faboideae are usually herbs or vines and often have once-compound leaves. The flowers are monosymmetric with the odd petal being adaxial and outside the others; the papilionoid flower consists of adaxial banner, paired wings and two connate petals forming the keel. The stamens have more or less connate filaments and the fruit is often explosively dehiscent, splitting doiwn both sides and the valves twisting; the seeds lack both a pleurogram and a linea fissura.

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. - 95/2830. World-wide, but esp. N. hemisphere.

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

"Spiraeoideae" + Amygdaloideae + Maloideae: sugar alcohol sorbitol, cyanogenic glycosides, flavones +, ellagic acid 0; G 5<, (1)2< ovules/carpel, placental obturator +.

Maloideae s.l. - Gymnosporangium rust common. - Predominantly temperate.

• Rosaceae are a heterogeneous family, but they may be recognised by their often compound, stipulate and serrate leaves that are never(?) 2-ranked. Their flowers have a valvate calyx, always free and often clearly clawed petals, usually many stamens, a well-developed nectary on the hypanthium or at the base of the stamens, and free carpels. There are often achenes or drupelets, but the whole fruit is fleshy in a variety of ways. Cyanogenesis is common (crush the leaves of Prunus, and smell), but there is more than one pathway involved.

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

BARBEYACEAE Rendle - inflorescence fasciculate, bracts and bracteoles 0; flowers small, hypanthium and nectary 0, P uniseriate; fruit indehiscent, P accrescent. - 1/1. N.E. Africa, Arabia.

• Barbeyaceae may be recognised by their opposite, entire, exstipulate leaves with dense white indumentum on the undersurface and their shortly pedunculate, axillary, cymose infloresences. The flowers are small, wind-pollinated and with a uniseriate perianth and the nutlets are enveloped by the accrescent perianth.

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; anthers long, extrose, opening from the apex; fruit beaked, segments opening adaxially, densely wooly inside, with columella. - 1/2. Socotra, Somalia.

• Dirachmaceae are shrubs that may be recognised by their small, strongly-toothed leaves which have subulate stipules that persist along the stem. The flowers have an epicalyx and are relatively large, with extrorse, oppositipetalous stamens the anthers of which clearly open from the apex. The capsules are densely wooly inside.

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

• Rhamnaceae are recognisable by their toothed, stipulate leaves that often have strong, parallel secondary veins and scalariform tertiary venation. Their flowers have stamens opposite the often clawed petals, a hypanthium with a nectary inside, and a valvate calyx that is ridged down the middle on the inner surface. The capsules are rather similar to those of Euphorbiaceae, the pericarp often separating into two layers and the endocarp twisting as dehiscence occurs. However, since the seeds are basal, there is usually no columella, while towards the base of the fruit there is often a conspicuous ring marking the place where the hypanthium was attached. In taxa such as Gouania with largely inferior ovaries the calyx, etc., is persistent. In lianes the shoots often develop somewhat laterally in the leaf axils and may have a bud at the very base.

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.

• Elaeagnaceae are often thorny and mostly deciduous shrubs that are recognisable by their spiral to opposite exstipulate leaves with entire margins; the plant is densely covered with lepidote indumentum, which often gives it a greyish or silvery appearance. The hypanthium is relatively long and there is no corolla. The fruit is invested by the accrescent hypanthium (cf. Nyctaginaceae), and so the fruit is like a berry or drupe.

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.

• Ulmaceae may be recognised by their usually 2-ranked, strongly serrate, stipulate leaves with asymmetrical bases; the pinnate secondary veins proceed to the apex of the teeth (not in Ampelocera). At least some of the small flowers are perfect and the fruits are flattened.

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.

• Cannabaceae may be recognised by their 2-ranked, serrate leaves with ascending veins that do not proceed straight into the teeth, more or less intrapetiolar stipules, and prominent prophyllar buds. The fruit is a rounded drupe. However, Humulus and Cannabis have secondary veins running into the teeth, and Humulus lacks buds at the base of the staminate inflorescence...

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

MORACEAE Link - Latex milky. - 38/1100. Mostly tropical to warm temperate.

• Moraceae are laticiferous plants with stipulate leaves and often boxy fine venation; the flowers are small and densely aggregated; the inflorescence axis is often large, becoming much accrescent and ± fleshy in fruit.

URTICACEAE Jussieu - A abruptly straightening, G pseudomonomerous, ovule basal, ± atropous. - 54/2625. World-wide, but mainly tropical.

• Urticaceae are variable in habit but have stipulate leaves with boxy venation. The small flowers are borne in branched, clearly cymose inflorescences - carpellate inflorescences may be capitate - and the anthers are often explosively dehiscent. Some genera related to Cecropia are trees with stout stems, deeply lobed or compound leaves, densely spicate inflorescences, and stamens that are not explosive.

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

CUCURBITALES Dumortier

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. - 7 families, 129 genera, 2295 species.

ANISOPHYLLEACEAE Ridley - K epidermis with mucilaginous inner walls, C open, ± enclosing groups of A, lobed or laciniate; embryo fusiform, largely hypocotylar, cotyledons at most small. - 4/34. Pantropical.

• Anisophylleaceae are a rather undistinguished-looking family. The plant may be anisophyllous, the leaves being rather coriaceous, often yellowish drying, entire, and often more or less asymmetrical at the base. The flowers are small and the ovary is inferior.

[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Begoniaceae + Datiscaceae + Tetramelaceae]]: uniseriate rays 0; filaments short 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 - 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 1 (or ?2), only one fertile. - 1/6. New Guinea to New Zealand, W. Pacific.

• Corynocarpaceae can be recognised by dissecting their flowers. These are quite distinctive, although small: the stamens are opposite the petals and alternate with fringed staminodes, and the ovary has a single style. Vegetatively they are rather less distinctive, with spirally-arranged, entire leaves with intrapetiolar stipules.

CORIARIACEAE Candolle - Leaves opposite; inflorescence racemose, bracteoles 0; C open, fleshy, 1 apotropous ovule/carpel, styles slender, stigmatic all around; fruit an achene, surrounded by accrescent C. - 1/5. Very disjunct: circum S. Pacific to China and Himalayas, Mediterranean.

• Coriariaceae can be recognised by their opposite, sessile, entire leaves with veins coming from the base and at most minute stipules; many branches look like compound leaves, apparently being of limited growth. The inflorescences are racemes and the flowers have a poorly differentiated, biseriate perianth and achenes that are surrounded by the accrescent corolla and have long, subbasal styles.

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 [plants dioecious?]; a roof over the ovary [= styles marginal], stigmas large, elongated, bilobed; placentation parietal, many ovules/carpel, stigmas elongated.

CUCURBITACEAE Jussieu - Tendrils +, bifid, lateral to the leaf; young stem with separate bicollateral vascular bundles; pericyclic sheath 0; cystoliths +; indumentum rough hairy/prickly, walls calcified; 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 - Often annuals; tendrils 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.

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

• Cucurbitaceae are easily recognised. They are vines or lianes with often rather roughly hairy exstipulate leaves that have palmate venation. In the leaf axil there is a usually lateral and branched tendril, a vegetative bud, a flower, or a yet more complex organisation. The plants are dioecious or monoecious, and the flowers have a hypanthium, a often complex androecium made up of twisted, monothecal anthers, and an inferior ovary with parietal placentation. The corolla is often more or less yellow, sometimes red. The seeds are more or less flattened.

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 - 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.

• Begoniaceae may be recognised by their succulent, herbaceous habit and often 2-ranked leaves the blades of which have asymmetrical bases and well developed teeth. The hairs are often large, flattened, or almost setular. The plants are monoecious, the staminate and carpellate flowers often have different numbers of tepals, both stigmas and anthers are bright yellow, and the fruit is often winged.

FAGALES Engler

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, 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.

• The family is readily recognisable in fruit. The distinctive cupules enclose one to three fruits, and the seed itself is often surrounded by dense hairs coming from the endocarp. The outer surface of the wood is characteristically furrowed, and the inflorescences are erect to pendant, spicate to catkinate. The leaves are usually spirally inserted, serrate (if entire the venation is camptodromous), and rather leathery and the hairs are stellate or branched.

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

[Myricaceae [Juglandaceae + Rhoipteleaceae]]: sieve tube P-protein bodies 0; peltate glandular hairs +; leaves spiral, stipules 0; one flower/bract; ovule single [per flower], atropous.

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

• Myricaceae are woody plants recognisable by the aromatic glands that cover the plant coupled with the spiral, simple, rarely deeply lobed leaves.

Juglandaceae + Rhoipteleaceae: leaves pinnate; endosperm 0; x = 16.

JUGLANDACEAE Perleb - Pericarp intrusive; seeds large, pachychalazal, cotyledons much folded. - 7-10/50. North temperate, S. to Argentina and Malesia.

1. Engelhardioideae Iljinskaya - Bracts 3-lobed; nuts with a fibrous layer. - 3-4/14. Himalayas to Malesia, Mexico to Colombia.

2. Juglandoideae Manning - Bracts unlobed; fruit with sclereids in shell. - 3-6/35. Temperate N. hemisphere, only 1 sp. in Europe, Central America and Andes.

• Juglandaceae may be recognised by the small, aromatic glands on the young parts of the plant; the compound, estipulate, usually spiral leaves the leaflets of which are often serrate; and the spicate inflorescences with single, axillary flowers. The endocarp of the drupaceous fruit has complex intrusions.

RHOIPTELEACEAE Handel-Mazzetti - Buds lacking scales; leaves 2-ranked, stipules +, asymmetrically caudate; 3 flowers/bract, central perfect; P 2 + 2, pollen colporate, G 2, ovules bitegmic, anatropous. - 1/1. China.

• Rhoipteleaceae may be recognised by their stipulate and odd-pinnately compound leaves with serrate leaflets, indumentum of peltate scales, and branched, pendulous inflorescences with flowers in triads, the central flower apparently being perfect and having a superior ovary.

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

CASUARINACEAE R. Brown - 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.

• Casuarinaceae are easily recognised, looking as they do rather like a conifer. The minute leaves are whorled, and the carpellate inflorescence forms a small cone. The bracteoles that gape widely at maturity to release the samara fruits are very distinctive.

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

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.

• Ticodendraceae are wind-pollinated trees that may be recognised by their doubly-serrate leaves that have stipules encircling the stem and carpellate flowers with two, long stigmas.

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

• Betulaceae are trees that are usually deciduous; they may be recognised by their stipulate, doubly serrate leaves, catkinate inflorescences, and fruits are that are either small and flattened or associated with large, leafy bracts.

EUROSID II: style single; endosperm scanty.

HUERTEALES Doweld

Vessel elements with scalariform perforations; flowers small, A = and opposite K, ovules 1-2/carpel; embryo at most medium. - 2 families, 4 genera, 21 species.

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

• Tapisciaceae can be recognised by their spiral, odd-pinnately compound or trifoliolate leaves with glands at the point of articulation of the petiolule with the rhachis; the leaflets have toothed margins.

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

• Dipentodontaceae can be recognised by their 2-ranked, serrate leaves and lobed stipules that are borne on the stem adjacent to the leaves. The simply umbellate inflorescences with small flowers and fruits that have almost indistinguishable and persistent sepals and petals are distinctive.

Brassicales + Malvales + Sapindales: endosperm scanty.

BRASSICALES Bromhead  

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, hypanthium +, C clawed, A 8, with short connective prolongation, placentation apical-axile, 1-2 epitropous ovules/carpel, style long.

AKANIACEAE Stapf - Leaves pinnate. - 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, C clawed, 2 tenuinucellate ovule/carpel. - 1/95. New World.

• Tropaeolaceae are rather fleshy vines with twining petioles. The broad leaf blades have palmate venation and may be palmately-lobed, palmate or peltate. The flowers are large, strongly monosymmetric, and have an adaxial calycine spur, five clawed petals, and eight stamens. The fruit is usually a three-lobed schizocarp, and each unit contains but a single seed.

Moringaceae + Caricaceae + Limnanthaceae + Setchellanthaceae + 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.

• Moringaceae are woody, often quite stout-stemmed shrubs or trees whose foliage often smells unpleasantly when crushed. They are recognisable by their spiral, deciduous, odd-pinnate, 1- to 3-compound leaves that have conspicuous glands at the articulations and at the base; often clearly obliquely monosymmetric and papilionoid flowers (the median petal is adaxial) with five stamens; and long, slender, tricarpellate, capsular fruits.

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).

• Caricaceae are stout-stemmed often prickly tree, rarely vines, with flowing, latex-like exudate and palmately-compound or -lobed leaves. The inflorescences are axillary, cymose, and the flowers - always imperfect - are moderate in size. The fruit is a berry and contains numerous seeds clearly borne on five parietal placentae and each surrounded by mucilage.

Setchellanthaceae + Limnanthaceae + Koeberliniaceae + Bataceae + Salvadoraceae + Emblingiaceae + Pentadiplandraceae + Gyrostemonaceae + Resedaceae + Tovariaceae + Cleomaceae + Capparaceae + Brassicaceae: nodes 1:1.

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

• Setchellanthaceae are pungent-smelling shrubs that may be recognised by their large, blue, axillary flowers with parts usually in sixes and an irregularly-splitting calyx. Vegetatively the plant is rather undistinguished, although it has T-shaped hairs and the rather small, entire, exstipulate leaves have secondary veins arising from near the base and lack much in the way of a petiole. The three-carpellate fruit is septifragal and there is a persistent columella.

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

LIMNANTHACEAE R. Brown - 1(2)/8. Temperate North America.

• Limnanthaceae are rather soft-stemmed herbs with spiral, exstipulate, deeply-lobed or compound leaves and rather open flowers with a valvate calyx, contorted corolla, and gynobasic style or styles. The fruit is a schizocarp separating into shortly spiny, one-seeded portions (mericarps).

Koeberliniaceae + Bataceae + Salvadoraceae + Emblingiaceae + Pentadiplandraceae + Gyrostemonaceae + Resedaceae + Tovariaceae + Cleomaceae + Capparaceae + Brassicaceae: style/stylodia short to absent; seeds exotegmic; embryo curved.

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

KOEBERLINIACEAE Engler - 1/1. C. and S.W. North America, Bolivia(!).

• Koeberliniaceae are thorny, much-branched shrubs of dry regions; the prophylls are basal on the branches/thorns. The flowers are small and undistinguished, and the fruit is a few(1-2)-seeded berry terminated by the persistent style.

Bataceae + Salvadoraceae: rays wide, multiseriate; nodes two trace; stomata paracytic; leaves opposite, with 2ndary veins ascending from at or near base; pollen psilate, 2 basal ovules/carpel; endosperm 0, embryo ± straight, color?

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

• Bataceae are halophytic subshrubs with opposite, small, entire, fleshy leaves that have minute "stipules". The flowers are staminate or carpellate and aggregated into dense, spicate, more or less strobilate, inflorescences. The carpellate flowers lack bracts or perianth and are fused together to form a compound fruit.

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

• Salvadoraceae are rather small-leaved shrubs, sometimes scrambling, or small treees with opposite, rather leathery leaves; small, 4-merous flowers with connate sepals and two carpels with a short style; and a fleshy fruit.

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.

• Emblingiaceae are low-growing herbs with axillary, ebracteolate and strongly monosymmetric flowers. The flowers are held upside down, the large, connate calyx is slit adaxially, there are two, abaxial, slipper-shaped petals (hence the specific epithet), and there is a curved androgynophore with an abaxially-positioned nectary at the base.

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; stylodia +.

GYROSTEMONACEAE Endlicher - G borne around the central axis in whorl(s). - 5/18+. Australia.

• Gyrostemonaceae are yellowish-drying (including the twigs!) shrubs with rather small, often quite narrow but otherwise featureless leaves. The small flowers with a persistent, connate calyx (presumably; there is only a single perianth whorl) and many stamens and/or carpels clearly arranged in a whorl are quite distinctive.

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.

• Resedaceae are yellowish-drying sometimes somewhat shrubby herbs with terminal, racemose inflorescences and rather small, monosymmetric flowers. The leaves are quite variable, although there are often small stipules. The petals are free and clawed; the adaxial petal is usually largest and the limb is often fringed. The extrastaminal disc/nectary is often conspicuous. The dry fruits often have an apical opening persisting between the stylodia (a quasi follicle) and the seeds are often arillate.

TOVARIACEAE Pax - 1/2. Tropical America.

• Tovariaceae are herbs that can be recognised by their trifoliate, stipulate leaves, terminal, racemose inflorescence, and 6-9-merous, polypetalous flowers with a short style and spreading stigmas.

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, persistent, woody placental strands well developed; seeds with invaginated coat.

CAPPARACEAE Pax - 16/480. Largely tropical.

• Capparaceae are woody, the inflorescence is racemose, the flowers have a gynophore and often many stamens with long filaments, and the fruit is indehiscent.

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.

• Cleomaceae are herbaceous or shrubby, the inflorescence is racemose, the flowers have four, clawed petals, six stamens, and two carpels; the gynoecium has a gynophore, the stamens have long filaments, and the dehiscent fruit has a persistent, woody placenta.

BRASSICACEAE Burnett - Methyl glucosinolates 0, erucic acid [fatty acid] +; roots lacking mycorrhizae; floral development closed, 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 +. - 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).

• Most Brassicaceae are herbs, and a number of taxa have distinctive small, branched, eglandular hairs. The ebracteate inflorescence is racemose, the flowers have four, clawed petals (the alternate name 'Cruciferae' for this family is taken from the cross-like arrangement of these four petals), six stamens with relatively short filaments (so the anthers are included), the gynoecium has two carpels and a short style, and the dehiscent fruit has both persistent, almost woody placental strands and a membranous false septum.

Malvales + Sapindales: ?

MALVALES Dumortier  

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

• Many members of this order can be recognised by their spiral, pulvinate leaves that often have palmate venation, or at least a pair of lateral veins leaving from the very base of the blade; stellate or fasciculate hairs and their derivatives are common. The bark is often very fibrous and tough because of the stratified phloem. The calyx is sometimes valvate, the corolla often contorted, the stamens usually numerous, the capsule sometimes muricate or spiny, and the seeds or inside of the carpel walls sometimes conspicuously hairy.

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.

• Neuradaceae are rather small herbaceous or subwoody plants of desert areas with small, toothed, exstipulate leaves and large flowers the petals of which dry a distinctive color; there are 5-10 separate styles.

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

THYMELAEACEAE Jussieu - Pericyclic fibers 0; crystal sand +; epidermal cells (massively) mucilaginous; stomata cyclocyclic. - 46/755. World-wide, esp. trop. Africa and Australia.

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

Octolepidoideae + Thymelaeoideae: C 0, 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; palisade endotegmen 0. - 37/690. World-wide, esp. trop. Africa and Australia.

• Thymelaeaceae are recognisable by their very fibrous bark and entire, exstipulate often opposite leaves which frequently have rather close, parallel venation and leave prominent raised scars on the stem after they fall off; the indumentum is of unicellular hairs and is often silky-adpressed. In many taxa, the twigs are notably flexible, and plants often smell unpleasantly. The flowers are quite often in heads, and the hypanthium is often long; any corolla lobes are usually at most small. The plants are often toxic.

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, micropyle bistomal.

SPHAEROSEPALACEAE (Warburg) Bullock - 2/18. Madagascar.

• Sphaerosepalaceae are a small family, but with a great deal of variation. The stipules are broad, intrapetiolar, and more or less encircling the stem. The flowers are usually four merous, and the sepals are imbricate in pairs; the fruits or carpels are single seeded.

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.

• Cistaceae are aromatic shrubs growing in open, sunny areas often on a sandy or chalky substrate. They often have opposite leaves with broad, even connate bases. The inner three K are sometimes described as being contorted, rather, the aestivation is extreme-quincuncial, the two outer sepals being much smaller than the others. The three inner sepals are contorted in the opposite direction to that of the petals. The free petals are crumpled when in bud and there are numerous stamens that are often sensitive to touch.

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

SARCOLAENACEAE Caruel - 8/60. Madagascar, mostly E. and C., fossil pollen in South Africa.

• The flowers, which have a massive "involucre" and sometimes also an enclosing bract (?bracteole), and an extrastaminal disc, are distinctive. The underside of the lamina of Sarcolaena has parallel lines. The hairs are stellate.

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

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

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

3. Dipterocarpoideae - 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.

• Dipterocarpaceae are trees that may be recognised by their often 2-ranked, coriaceous leaves with strong, parallel, if not particularly close, secondary veins and scalariform tertiaries; the hairs are often fasciculate or stellate. The inflorescences are often distinctively monochasial, and the small flowers are pointed in bud and have a conspicuously contorted cotrolla. Fruits with strongly accrescent but often unequal sepals surrounding single-seeded nuts are distinctive and common in the family, but there are other fruit types.

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

• Muntingiaceae can be recognised by their 2-ranked, toothed, leaf blades with asymmetrical bases and heteromorphic stipule-like prophylls. Their flowers are in extra-axillary fascicles, the calyx is valvate, the corolla shortly clawed and crumpled in bud, and the stamens are numerous.

MALVACEAE Jussieu - 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.

• Malvaceae are usually readily recognisable even when sterile by the combination of fibrous bark, alternate, stipulate leaves with toothed margins, ± palmate venation, and stellate to lepidote indumentum. Mucilage is common (cf. Elaeocarpaceae!). The flowers and fruits are also distinguishable by a combination of characters. The flowers often have a valvate, connate calyx with a nectary at the base inside and a contorted corolla; the stamens are often numerous and variously connate and/or fasciculate. The inside wall of the fruit and/or the surface of the seed is often hairy.

CYTINACEAE A. Richard - Echlorophyllous root parasites; vessels 0; 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.

• Cytinaceae are echlorophyllous parasites with racemose inflorescences, sometimes capitate, in which the individual flowers are moderately sized and easy to make out. The more or less spreading perianth is a single whorl, basaly connate, and the stamens are extrorse (staminate flowers) and the style long and expanded like a round disc at the apex (carpellate flowers).

SAPINDALES Dumortier  

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

BIEBERSTEINIACEAE Endlicher - 1/5. Greece to Central Asia.

• Biebersteiniaceae are perennial, glandular-hairy herbs that have odd-pinnate leaves with lobed to compound leaflets and petiolar stipules. Ihe inflorescence is erect and racemose and the flowers are large. The fruit has a single seed per carpel and a persistent columella.

KIRKIACEAE (Engler) Takhtajan - Inflorescence dichasial, flowers small, 4-merous; fruit a schizocarp, columella persistent. - 2/6. Tropical and S. Africa.

• Kirkiaceae are woody plants that have often rather closely-set odd-pinnate leaves with toothed leaflets. Ihe inflorescence is dichasial and the flowers are small and 4-merous. The fruit is a ridged or angled schizocarp with persistent if slender columella strands.

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.

• Rutaceae are recognisable because of their often opposite, compound and punctate leaves; their cymose inflorescences; their flowers, with the stamens often arranged in a ring, thick filaments, conspicuous disc at the base of the ovary, and expanded stigma, are quite distinctive. Although the fruits are variable, many have a glandular-punctate pericarp, while others are dry, deeply lobed, and dehisce to reveal the black and shiny seeds.

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

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

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

• Meliaceae are recognisable by their usually spiral and odd-compound leaves with well-developed leaf buttresses, i.e. the leaf bases are swollen and more or less elongated vertically. The leaflets are often opposite and dry thin and brittle, with little in the way of tertiary venation evident. The bark may smell - sweet, like garlic, etc. - and milky exudates occur sporadically. The flower buds are often longer than broad, with petals much longer than the sepals, and most members have a staminal tube and a large stigma. The fruits and seeds are usually quite large.

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

• Simaroubaceae can be recognised because the twigs have conspicuous pith - herbarium specimens are often notably light - and the bark is bitter, even when dry; the leaflets of the odd-compound leaves often have rather coarse, blunt, teeth and flat glands on the lower surface, sometimes close to the margin, and the rhachis collapses at the nodes. The flowers are rather small, the petals usually barely exceeding the sepals. The fruits are often separate drupelets or samaras.

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.

• Anacardiaceae may be fairly readily recognised because of their often black and/or rather resinous-smelling exudate; the leaves often have black discolorations and the blade may dry a distinctive grey colour. In herbarium specimens, the dry, black exudate may entirely cover the cut surface of the twig, or it may form a distinctive black ring around the periphery. The leaves are often odd-pinnate and the leaflets are opposite to alternate. The flowers are small, and the fruits are often flattened, operculate drupes with excentric style or styles.

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

1. Beiselia - Vessel elements with scalariform perforations; carpels 9-12, ovules superposed. - 1/1. Mexico.

The rest - Cotyledons folded, palmately lobed. - 17/550. Tropical, but esp. America and N.E. Africa.

• Burseraceae are recognisable by their often scented resinous exdate; compound, odd-pinnate leaves with opposite, often long-petiolulate and more or less pulvinate leaflets that have prominent fine venation especially when dry; and petioles, petiolules and rachis that tend to be brownish and more or less scurfy. The swollen leaf base is often concave adaxially. The calyx is often connate and 3- or 4-lobed, and the angled drupes have a wrinkled surface when dry. The resinous exudate often smells of almonds (myrrh, frankincense), and in some genera the leaflets are strikingly symmetrical.

SAPINDACEAE Jussieu - Cyclopropane amino acids +; disc outside A, A 8 or fewer, ovules sessile; embryo curved, the radicle in a pocket of the testa. - 135/1580. ± World-wide.

1. Xanthoceras - Leaves deciduous; disc with golden, horn-like glands, 7-8 ovules/carpel. - 1/1. N. China.

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

3. Dodonaeoideae - 2 basal apotropous (1 apical epitropous) ovules/carpel. - Pantropical-warm temperate, esp. Australia/Southeast Asia.

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

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.

• Martyniaceae are herbs with sticky hairs and large, monosymmetric flowers with a superior ovary. The placentation is parietal, and the fruits have two prominent apical hooks as well as other hooks, spines or prickles.

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

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

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, A 4 or 2 + 2 staminodes, the abaxial members Z-shaped, with an appendage, or much reduced, stigma bilobed, sensitive; seeds with ruminate endosperm. - ca. 6/170. Pantropical to warm temperate.

• Many Linderniaceae are rather small herbs with opposite leaves, square stems, and monosymmetric flowers with a sensitive stigma and the lower stamens usually clearly different from the upper, either Z-shaped or long and curved, with a projection, or simply very much reduced.

BIGNONIACEAE Jussieu - Leaves bicompound; tapetum amoeboid, pollen colpate, bundles in the ovary wall and also opposite septum, 2 series of ovules/carpel; 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.

• Bignoniaceae can be recognised quite easily. Vegetatively, they are woody and with lenticillate stems, and most have opposite, compound leaves; the vines and lianes often have leaf tendrils. The often rather large flowers are monosymmetric, and there are often nectaries on the leaf, stem (at the nodes), calyx and even the ovary surface. The calyx is often spathaceous (Old World taxa), there are two series of ovules in each carpel, and the broad, almost spathulate stigmatic lobes are often sensitive. The seeds are usually flattened and winged.

VERBENACEAE Jaume Saint-Hilaire - Inflorescence racemose; G with two ovules/carpel, stigmatic head swollen. - 30/1100. Pantropical (to warm temperate), but mostly New World.

• Verbenaceae are shrubs or trees that may be recognised easily because the plants are often aromatic, the opposite leaves are serrate, the stem is often angled and with a line across the node, and the monosymmetric flowers are borne in racemes or heads. The stigmatic area is conspicuously swollen and glandular and the ovary has four ovules.

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.

• Lamiaceae are herbs, also shrubs and lianes, that may be recognised by their opposite, usually serrate leaves, more or less square stems, and monosymmetric flowers borne in often clearly cymose clusters. The plants are often aromatic. The often gynobasic style is apically bifid, the two lobes being sharply pointed, and the four ovules of the bicarpellate gynoecium are borne in uniovular compartments. The calyx is an integral part of the dispersal mechanism and is or more or less accrescent; it may be fleshy, or distinctively ridged and dry. Woody members often have strongly lenticillate twigs.

SCHLEGELIACEAE (A. H. Gentry) Reveal - Fruit baccate. - 4/25. Mexico to tropical South America, Cuba (Synapsis).

• Schlegeliaceae may be recognised by their woody habit, white bark, opposite, often thick leaves at least sometimes with small surface glands on the lower surface, monosymmetric flowers, and berries.

PHRYMACEAE Schauer - Ca 19/234. ± World-wide, esp. temperate and W. North America and Australia, but few humid tropics.

ACANTHACEAE Jussieu - Acicular fibers +; endosperm development asymmetrical [the two haustoria come to lie close to each other]. - 229/3500. Mostly tropical.

1. Nelsonioideae (Nees) Sreemadhavan - 6-7/170. Tropical.

Acanthoideae + Thunbergioideae + "Avicennioideae": acicular fibers +; (inverted vascular bundles in the pith); (abaxial C lobes outside others), pollen other than colporate, 2-few collateral ovules/carpel, endothelium 0; endosperm 0.

2. Acanthoideae - Capsules explosive, obovoid, seeds borne on hook-like hardened funicles. - 217/3220. World-wide; the bulk of the family.

Thunbergioideae + Avicennioideae: ovules 2/carpel; 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, or up to 16 lobes, 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.

• Acanthoideae are usually herbs, and can be recognised by their often swollen nodes, the stem immediately above them collapsing on drying, and the opposite leaves that often have cystoliths (but only in Acanthoideae, and even then not Acanthus and immediate relatives!). The bracteoles are often prominent and the calyx (and corolla) lobes may be narrow. The obovoid, almost woody capsules have few, relatively large, flattened seeds cradled by hook-like jaculators; the latter are hardened funicles.

• Thunbergioideae are vines. There are no bracts, but large bracteoles envelop the flower. The calyx is reduced to a rim or is represented by a number of small lobes. The flowers are large, the anther connective is elongated and there are two ovules per carpel.

• Avicennioideae are mangrove trees with pneumatophores that can be recognised by their opposite, entire, somewhat fleshy leaves with invisible secondary and finer venation, clavate hairs covering the lower surface, and salt glands on both surfaces. The flowers have as many stamens as corolla lobes, a persistent green calyx, and achenes with a single rather large seed.

• Nelsonioideae are rather undistinguished!

SCROPHULARIACEAE de Jussieu - 65/1700. World wide.

OROBANCHACEAE Ventenat - Stomata permanently open; glandular hairs lacking vertical walls in the heads; inflorescence racemose; abaxial lobes of C often outside others in bud [aestivation descending cocleate]. - 99/2061. World wide, but especially N. (warm) temperate and Africa-Madagascar.

• Orobanchaceae are herbs, rarely shrubs, mostly with racemose inflorescences. In the flower, the abaxial lobes of the corolla are outside the others in bud. Nearly all members of the family have a holo- or hemiparasitic life-style. Some Plantaginaceae have similar corolla aestivation, but they are autotrophic and have glandular hairs with the heads divided by vertical cell walls.

PAULOWNIACEAE Nakai - 1/6. (Warm) temperate East Asia.

• Paulowniaceae are trees that may be recognised by their opposite leaves and terminal inflorescences with large, monosymmetric flowers; the calyx is densely covered with brown indumentum.

BYBLIDACEAE (Engler & Gilg) Domin - Rhizomatous herbs; 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.

• Byblidaceae are rhizomatous herbs with linear leaves that have densely covered by glands. The larger glands, with their heads formed from a single layer of cells radially divided, are conspicuous and look like little parasols; the sessile glands are smaller versions of these. The axillary, polysymmetric flowers have a broadly-spreading, contorted corolla that is only shortly connate at the base, anthers that are porose and whose apices form a cone, and a long style.

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.

• Lentibulariaceae are vegetatively diverse insectivorous herbs of wet habitats. Pinguicula has flat leaves that are sticky on the adaxial surface, Genlisea has tubular leaves and forked, subsurface traps with the opening spiralling along the branches of the fork, while Utricularia may have leaves, but the rest of the vegetative body is impossible to equate with conventional plant parts and there are little insect-collecting bladders with trap-door entrances that open on stimulation of one of the four sensitive hairs. The flowers are much more uniform. They are strongly bilabiate, with a nectar-secreting spur, and there are only two anthers.

SOLANALES Dumortier  

Alkaloids, O-methyl flavonols +; corolla plicate, 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; A extrorse, G inferior, style short, stout; seeds winged. - 3/5. Africa and Madagascar.

• Montiniaceae are woody plants that can be recognised by the persistent short style with large stigmatic lobes that caps the smooth fruit. The often 4-merous flowers are small, with free, valvate petals and extrorse, more or less basifixed anthers. Crushed tissue may have a spicy or acrid smell.

Sphenocleaceae + Hydroleaceae: placentae swollen [?level]; endosperm cellular, at most scanty, with multicellular micropylar and chalazal haustoria.

SPHENOCLEACEAE (Lindley) Baskerville - Rather fleshy annual herbs; fructose +; stomata tetracytic; inflorescences spicate; disc 0, G ± inferior, style short; fruit capsular, circumscissile. - 1/2. Pantropical.

• Sphenocleaceae are erect fleshy annuals of damp or flooded habitats that have dense spikes of small flowers with more or less inferior ovaries. The small fruits are circumscissile, the lid coming off just below the point of origin of the calyx. Older stems have a green reticulation marking the plates of cells bounding the cortical air spaces.

HYDROLEACEAE Berchtold & J. Presl - 1/12. Tropical.

Convolvulaceae + Solanaceae: caffeic acid esters, tropane [polyhydroxynortropanes] and pyrrolidine alkaloids, acylated anthocyanins +; internal phloem +; C usu. contorted or induplicate-valvate; K persistent 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.

• Convolvulaceae are recognisable by their usually viny habit and leaves with palmate venation; the plants are often obviously laticiferous. The quincuncial calyx is often notably persistent; the sepals are free, and are rarely green in fruit, often being brown and woody - as in wood roses (Merremia tuberosa) - to scarious. The large, radially symmetric corolla has contorted or induplicate-valvate aestivation and the five stamens are adnate to the tube. The gynoecium is bicarpellate, each loculus usually having two ovules (you rarely need to check this since the calyx is so distinctive), and the seeds have a notably hard, shiny testa.

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 - 4/7. Greater Antilles, but not Jamaica, South America.

4. Cestroideae - 8/195. South and Central (and North) America.

5. Petunioideae - 13/160. Central and South America.

Solanoideae + Nicotianoideae: x = 12.

6. Solanoideae Schlechtendal - Leaves often geminate and/or branching apparently not axillary; fruit a berry; seeds flattened. - 61/1925. World-wide, but esp. South America and others N. temperate.

7. Nicotianoideae - 8/125. Mostly Australian, also North and South America, Africa.

• Solanaceae are quite often rather foetid herbs or weak shrubs with polysymmetric or weakly bisymmetric sympetalous flowers - when bisymmetrical, the flowers are held so that "the" odd petal is in the adaxial position. Vegetatively many Solanaceae - especially Solanoideae, the bulk of the family - are distinctive because the insertion of leaves, branches and flowers often does not seem to follow any regular sequence; paired leaves adjacent on the stem are common. The flowers often have porose anthers or the anthers as a group are connivent at the apex. The fruit is usually bicarpellate and is a septicidal capsule or a berry with flattened seeds; the calyx consists of connate sepals that persist and even enlarge in fruit.

EUASTERID II: vessel elements with scalariform perforations; flowers rather small, style short; endosperm copious, embryo short/very short.

AQUIFOLIALES Senft

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.

• Cardiopteridaceae are a rather heterogeneous group of genera, although one or sometimes two rather slender styles are usually evident. The fruits are more or less flattened and/or ridged drupes.

STEMONURACEAE (M. Roemer) Kårehed - 12/80. Tropics, esp. Indo-Malesia to Australia (Queensland).

• Stemonuraceae are woody plants whose entire, exstipulate leaves lack much in the way of venation when dry. The flowers are rather small and are borne in cymose inflorescences; the stout filaments are often conspicuously hairy, the broad stigma is sessile, and the gynoecium is pseudomonomerous. The fruit is a more or less flattened and ridged drupe with one side sometimes very different in appearance, both in color and texture, from the other, as in Medusanthera.

Phyllonomaceae + Helwingiaceae + Aquifoliaceae: nodes 1:1; leaf margins toothed, stipules small, cauline; drupe with separate pyrenes.

PHYLLONOMACEAE Small - Petiole bundle annular; G inferior, several ovules/loculus; fruit a berry. - 1/4. Mexico to Peru.

• Phyllonomaceae are shrubs or small trees that may be recognised by their leaves which have epiphyllous inflorescences borne on the upper surface of the lamina towards the apex; the stipules are minute. The ovary is inferior and has a prominent disc; the fruit is fleshy and contains only a few seeds.

Helwingiaceae + Aquifoliaceae: ?

HELLWINGIACEAE Decaisne - P 3-5, A = and alternating with P, pollen with diffuse endoapertures. - 1/3. Himalayas to Japan.

• Helwingiaceae are evergreen shrubs or small trees that may be recognised by their leaves which have epiphyllous fasciculate inflorescences on the upper surface of the blade; the stipules are small and fimbriate. The flowers have only a single perianth whorl, the ovary is superior, and the fruit is a drupe.

AQUIFOLIACEAE A. Richard - 1/405. ± World-wide, esp. America and South East Asia-Malesia, few in Africa.

• The family may be recognised by its usually spiral, toothed leaves which may have small cork-warts on the lower surface of the lamina; the stipules are minute and blackish. The inflorescence is cymose and frequently subumbellate. The fruits often dry broadly bottle-shaped, being capped by the distinctive, sessile, broad, black stigma; there are several stones.

ASTERALES + APIALES + DIPSACALES: iridoids +; C tube initiation early, G [2-3], inferior.

Unplaced:

BRUNIACEAE Berchtold & J. Presl - 12/75. South Africa, almost entirely the Cape Province.

• Bruniaceae are ericoid shrubs from the Cape that can be recognised by the often black projection at the apex of the leaf blade and the often small flowers aggregated into more or less capitate inflorescences. The ovary is more or less inferior.

ASTERALES Lindley  

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 Candolle - 4/13. Mauritius, New Guinea to New Zealand.

1. Rousseaoideae - 1/1. Mauritius.

2. Carpodetoideae (Fenzl) J. Lundberg - 3/12. New Zealand, E. Australia, Papuasia.

• Rousseaoideae may be recognised by their whorled, serrate leaves, bud scales, and their single, terminal flower with large calyx that persists reflexed on the fruit, long sagittate anthers, and apparently superior ovary tapering gradually into a stout, persistent style.

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 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 Lammers - C of 3 adaxial and 2 abaxial lobes, 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.

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 (Jussieu) 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 (A. DC.) Schönland - Roots tuberous; A basally connate, 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]. - 1/50. Africa, esp. the south, Cape Verde Islands.

• Campanulaceae are usually herbs that can be recognised by their white latex, often rather soft leaves, and flowers with an inferior ovary and secondary pollen presentation.

• Polysymmetric campanulate flowers with sprawling dehisced anthers at the base inside distinguish Campanuloideae; pollen is held among hairs along the style, and these hairs later retract. There are often three carpels.

• Lobelioideae have monosymmetric flowers in which the corolla may be split to the base down one side and the anthers (and filaments) are connate and the former at least initially enclose the stigma. The style pushes up within the anther tube and exposes the pollen. There are two carpels.

• Nemacladoideae have monosymmetric flowers in which the filaments are more or less connate and the anthers are free, flipping backwards after they have dehisced. There are two carpels.

• Cyphioideae also have monosymmetric flowers in which the stamens are more or less completely free. There are two carpels.

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.

• Pentaphragmataceae are distinctive plants. They are rather fleshy herbs with 2-ranked, asymmetrical, usually serrate leaf blades and scorpioid cymes with sessile flowers. The flowers are radially symmetrical and have relatively large, conspicuous sepals, extrorse anthers and an inferior ovary; there are nectariferous pockets between the septae that join the hypanthium to the ovary. The fruit is baccate.

Alseuosmiaceae + Phellinaceae + Argophyllaceae + Stylidiaceae + Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae: ?

Alseuosmiaceae + Phellinaceae + Argophyllaceae: 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.

• Alseuosmiaceae may be recognised even vegetatively by their spiral, serrate leaves with axillary tufts of uniseriate hairs. Their flowers have an inferior ovary and usually a tubular corolla, and the corolla lobes have fringed or erose winged margins. The fruit is baccate and contains rather small seeds.

Phellinaceae + Argophyllaceae: pollen (spiny) with rugulose exine, ovules apotropous, style short.

PHELLINACEAE (Loesener) Takhtajan - 1/12. New Caledonia.

ARGOPHYLLACEAE (Engler) Takhtajan - 2/17 S.W. Pacific.

STYLIDIACEAE R. Brown - A extrorse, 2-3, pollen colpate. - 6/245. South East Asia to New Zealand, S. South America.

1. Donatioideae B. Chandler - 1/2. New Zealand, Tasmania, S. South America.

2. Stylidioideae - Flowers semi-resupinate, split-monosymmetric, A 2, adnate to style. - 5/155. South East Asia, Malesia, S. South America, esp. Australia.

• Stylidiaceae are usually rosette or tussock herbs with sessile or obscurely petiolate leaves. Stylidioideae have very distinctive semi-resupinate, split-monosymmetric flowers. The two stamens are adnate to the style, the extrorse anthers being borne near the stigma, and the whole complex is often sensitive, moving when brushed by pollinator (hence the name, trigger plants). Donatioideae can be recognised by their solitary, polysymmetric, terminal flowers that have a variable number of free sepals and petals and two to three stamens with extrorse anthers.

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; corolla lobes with marginal wings or fringes, G superior, placentation parietal. - 5/ca 40. World-wide.

• Menyanthaceae are aquatic plants with orbicular to palmately-compound leaves with broad bases and quite large usually heterostylous polysymmetric flowers. The petals are connate, the corolla lobes have disinctive marginal wings or fringes and the ovary is superior

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 - Stylar cup. - 12/440. Largely Australian, Scaevola pantropical.

• Goodeniaceae are more or less herbaceous plants with spiral leaves and monosymmetric flowers. The corolla is often divided to the base adaxially, the lobes have marginal wings that may make lobe each look trilobed, and the style is curved, with an apical cup and (in older flowers) a bilobed stigma in the middle. The anthers dehisce in bud and the ovary is inferior.

Calyceraceae + Asteraceae: flowers small, C tubular, commissural veins connate (median veins 0), filament collar +, G [2], with 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.

• Calyceraceae are herbs that may be recognised by their capitate inflorescences the flowers of which often open centripetally. The flowers are small and polysymmetric, the sepals are spines or are thick and aerenchymatous, the outer layer of the tubular corolla is photosynthetic, the stamens are free, and the ovary is inferior.

ASTERACEAE Martynov - Sesquiterpene lactones, terpenoid essential oils, iridoids 0; inflorescence capitulate, involucrate, ebracteate; K reduced, anthers connate, endothecial cells elongated parallel to main axis of anthers; tapetum amoeboid, ovule basal, epitropous. - 1620/22750. World-wide.

1. Barnadesioideae (Bentham and J. D. Hooker) Bremer & Jansen - pollen with intercolpar depressions. - 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.

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 +.

5. Carduoideae Kitamura - 83/2500. World-wide, but most N. hemisphere, esp. Eurasia/N. Africa.

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: deletion in ndhF gene.

8. Gymnarrhenoideae Panero & Funk - Plant amphicarpic; heads grouped into synflorescence; pappus of bristles. - 1/1. North Africa to the Middle East.

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.

• Asteraceae are very variable vegetatively, but may be recognised by their capitulate and involucrate inflorescences in which the numerous small centripetally-opening flowers are only sometimes subtended by bracts. The anthers are usually connate, forming a tube through which the stigma extends before the two lobes separate and become recurved.

Unplaced: There appear to be a number of unplaced taxa, many ex Saxifragaceae s.l., that are at the base of Apiales, perhaps.

Polyosmaceae + Tribelaceae + Eremosynaceae + Escalloniaceae: cork superficial; nodes 1:1; petals free.

POLYOSMACEAE Blume - 1/60. E. Himalayas and S. China to N.E. Australia and New Caledonia.

• Polyosmaceae are trees with opposite, often serrate, exstipulate leaves and axillary, racemose inflorescence. The flowers are 4-merous, with long buds, rather narrow petals, and an inferior ovary.

ESCALLONIACEAE Dumortier - cells in heads of gland hairs with vertically radiating walls; petiole bundle arcuate. - 8/68. Scattered: Central and South America, Malesia to New Zealand, Réunion.

Paracryphiaceae + Quintiniaceae + Sphenostemonaceae: true tracheids +; nodes 3:3; leaves spiral, serrate; inflorescence terminal, racemose; flowers 4-merous; capsule septicidal.

PARACRYPHIACEAE Airy-Shaw - P decussate-cochleate, caducous, A 2xP, filaments stout, thecae ± embedded in connective, disc 0, G [8-15], attached to central tissue, 4 crassinucellate ovules/carpel; fruit with carpels pulling away acropetally and opening adaxially, columella persistent. - 1/1. New Caledonia.

QUINTINIACEAE Dumortier - Peltate glands +; pollen 4-6 colporate. - 1/25. Philippines and New Guinea to New Zealand and New Caledonia.

• Quintiniaceae are evergreen trees or shrubs with spiral, serrate, estipulate leaves; the plant is covered with peltate scales. The inflorescence is terminal and the flowers have free sepals and petals and an inferior ovary. The fruit is a septicidal capsule.

SPHENOSTEMONACEAE P. van Royen & Airy Shaw - 1(2) pendulous ovule/carpel, funicular obturator +, style 0, stigma large, capitate; fruit a drupe. - 1/10. S. W. Pacific.

• Sphenostemonaceae are usually small, evergreen trees with spiral, strongly serrate, estipulate leaves and terminal inflorescences of rather small flowers. The sepals and petals are free, the anther thecae are borne on stout filaments, and there is no style; the fruit is fleshy and two-seeded.

APIALES Nakai

Group II decarboxylated iridoids +; ?nodes; ?stomata; inflorescence terminal, paniculate; plants dioecious; pedicels articulated; flowers 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 - 1/4. E. Australia, New Zealand.

Torricelliaceae + Griseliniaceae + Araliaceae + Pittosporaceae + 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 (Wangenheim) H. H. Hu - 3/10. Madagascar, South East Asia and W. Malesia.

Griseliniaceae + Araliaceae + Pittosporaceae + 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.

Araliaceae + Pittosporaceae + 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]; K small, G [2], both fertile, with nectariferous stylopodium, 2 epitropous ovules/carpel; fruits ± laterally compressed; hemicellulosic seed reserves common; x = 12.

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.

• Araliaceae may be recognised quite easily. They are often rather stout-stemmed and little-branched shrubs or trees, rarely herbs, with large and prominent scars from the fallen leaves; the plant may smell strongly. The leaves are often variously compound and with broad bases, although these latter rarely more or less encircle the stem; stipules are common and are morphologically diverse, although they are generally borne on the leaf base. Petioles are often long, and/or they vary considerably in length on the one shoot. The ultimate units of the inflorescence are umbels or heads, the flowers are often rather small, there are often three or more carpels and the petals are valvate; the fruit is drupaceous, sometimes flattened laterally, and the seeds may be ruminate.

Pittosporaceae + Myodocarpaceae + Apiaceae: ?

Pittosporaceae + Myodocarpaceae: ?

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.

• Pittosporaceae are aromatic woody and sometimes thorny plants with simple, estipulate leaves and rather conspicuous flowers that have sepals, petals and stamens equal in number and a bicarpellate gynoecium with parietal placentation. The fruits are often capsular and enclose pulpy-resinous seeds; the calyx is deciduous.

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 - Leaves often broad, with hairy or thorny tips to the teeth; fruit scaly or spiny. - 11/335. 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.

• Apiaceae are recognisable by their usually herbaceous habit, compound leaves with broad, sheathing base, inflorescences that are usually umbels of umbels, rarely heads, and polypetalous flowers usually with a minute calyx, clawed petals with incurved apices, and two carpels; the fruits are dry and schizocarpic.

• Saniculoideae have usually undivided leaves with hairs or spiny tips to the teeth, sessile spiny or scaly fruits, an often more or less simple umbel or head, and there is no free carpophore.

• Apioideae usually have more or less divided leaves, compound umbels, and the fruit has a free, bifid carpophore.

Unplaced: sister to Dipsacales, perhaps, in which case the characters of the order will need considerable modification....

Columelliaceae + Desfontainiaceae [= Columelliaceae s.l.]: 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, anthers basifixed, connective well-developed, style long, many ovules/carpel; K persistent in fruit; seeds many, anticlinal walls of exotestal cells thickened, with plasmodesmata.

COLUMELLIACEAE D. Don - A 2, latrorse, thecae semicircular, attached their length to expanded connective, stigma with two broad lobes, almost cup-like; fruit a septicidal and part loculicidal capsule. - 1/4. Colombia to Bolivia, in the Andes.

• Columelliaceae are evergreen shrubs with opposite, serrate leaves. Their flowers are distinctive, having two stamens with latrorse, semicircular thecae attached their length to an expanded connective, and an almost cup-like stigma with two broad lobes.

DESFONTAINIACEAE Endlicher - Leaf teeth spiny; flowers single; C contorted; anther thecae ± embedded in connective, G [5(-7)]; fruit a berry; outer walls of testa pectic; endosperm starchy. - 1/1. Costa Rica to Chile.

• Desfontainiaceae are evergreen shrubs that may be recognised by their opposite spiny leaves, single terminal flowers that have a long-tubular corolla, stamens borne in the throat and contorted lobes. The fruit is a berry with a persistent calyx.

DIPSACALES Dumortier

Group I secoiridoids +; nodes 3:3; buds with scales; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence terminal, cymose; K persistent in fruit; testa vascularised, exotestal cells enlarged, variously thickened and lignified. - 2 (7) families, 45 genera, 1090 species.

ADOXACEAE Trautvetter - C rotate. - 5/200. North temperate, tropical montane, but not in Africa.

1. Viburnum - Hairs stellate and variants. - 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.

• Adoxaceae may be recognised by their opposite, toothed leaves that are compound or simple. The inflorescence is usually flat-topped, cymose, and has numerous small flowers. The flowers are 5- or sometimes 4-merous, the calyx is open during development, the corolla is rotate, the style is at most short, and the fruit is drupaceous. Some Cornaceae look rather similar to Viburnum, but the former often have 4-merous flowers, they lack stellate indumentum, and their lateral veins ascend towards the apex of the blade.

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; 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 + Dipsacaceae + Valerianaceae: 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, G with sterile loculi much reduced, 1 apical ovule/flower, prominent endothelium with crystal layer; exotesta not thickened; endosperm scanty, embryo large; distinctive expansion of the chloroplast inverted repeat.

MORINACEAE Rafinesque - Ethereal oils +, iridoids 0; epicalyx 12-ribbed. - 2/13. Balkans to China.

• Morinaceae are robust, perennial herbs. They may be recognised by their opposite leaves with connate bases and often spiny margins and inflorescences consisting of whorls of flowers, each flower having an epicalyx. The flowers are sympetalous and strongly monosymmetric.

Dipsacaceae + Valerianaceae: saccharose +, quercetin 0; flowers rather small, A ± equal in length, tapetal cells 4 or more nucleate [?level], exine columellae prominent and branched; embryo green; chromosomes "large".

DIPSACACEAE Jussieu - Starch almost 0; inflorescence capitate, epicalyx 8-ribbed; tapetum glandular, micropyle very long; endosperm +. - 11/290. Eurasia, Africa, esp. Mediterranean region.

• Dipsacaceae are herbs with opposite leaves, capitate, involucrate inflorescences and more or less bisymmetric flowers each with a well-developed epicalyx. The dry fruit has a single seed.

VALERIANACEAE Batsch - Foetid monoterpenoids and sesquiterpenoids, valpotriates +; bracteoles +, epicalyx 0; pollen colpate. - 17/315. N. temperate, Andean South America.

• Valerianaceae are often foetid herbs with opposite, sometimes connate leaves and rather small flowers. The inflorescence is cymose and bracteolate. The sympetalous corolla is often spurred but other than that may be only weakly monosymmetric; the calyx often develops greatly in fruit.

UNPLACED EUDICOTS

APODANTHACEAE (R. Brown) 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.

HAPTANTHACEAE C. Nelson - Nectary 0; staminate flowers: P?, A 2, endothecium massive; carpellate flowers: P ca 4, G [3], placentation parietal. - 1/1. C. America, Honduras.

HOPLESTIGMATACEAE Engler & Gilg - 1/2. W. Africa.

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.

• Medusandraceae are trees whose stipulate leaves have long petioles that are swollen at the apex and weakly serrate blades with strong veins arising from at or near the base. The axillary inflorescences are almost catkin-like, the flowers having long and conspicuous staminodes borne opposite the sepals. The fruit is single seeded and capsular, and the sepals are much accrescent, becoming strongly recurved.

METTENUISACEAE Schnizlein - 1/7. Panama and N.W. South America.

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