EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, axillary buds +[?], prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable; P not sharply differentiated, outer members not enclosing the rest of the bud, smaller than inner members; A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, nucellus at apex of ovule 1-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; pollen germinating in less than 3 hours, siphonogamy, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm; tube moves between nucellar cells, double fertilisation +, endosperm diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA + C/PHYB + E gene pairs.

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous... For other features such a a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), where on the tree a thicker nucellus and a stylar epidermal layer are acquired has not yet been indicated.

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

EUDICOTS: myricetin, delphinidin scattered, asarone 0 [unknown in some groups, + in some asterids]; root epidermis derived from root cap [?Buxaceae, etc.]; nodes 3:3; stomata anomocytic; flowers (dimerous), cyclic; K/outer P members with three traces, "C" with a single trace; A few, (polyandry widespread, from few initial [5, 10, ring] primordia), filaments fairly slender, anthers basifixed; microsporogenesis simultaneous, microspore walls developing by centripetal furrowing; pollen with endexine, tricolpate; G with complete postgenital fusion, stylulus/style solid [?here]; seed coat?

THIS LOT (but getting fewer and fewer) UNPLACED  Main Tree, Synapomorphies.

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

Evolution. Wikström et al. (2001) suggest a stem group age of 75-72 million years before present, with crown group divergence beginning 49-46 million years before present.

Members of this group are not often eaten by butterfly larvae, but Lycaenidae-Riodininae-Hamearini and a few Riodinini (see also Abisara) are found on them, especially on Maesaceae but not so far on Samolus and Theophrastaceae (Ehrlich & Raven 1964).

Chemistry, Morphology, etc. Leaves of Theophrastaceae and Myrsinaceae are often described as being involute (?supervolute, cf. Cullen 1978) or conduplicate. There are common stamen/corolla primordia born on a ring primordium in this clade. but there is variation in the position/relative development of these primordia. In some cases such as Cyclamen the stamens are initiated as adaxial outgrowths of a common primordium, i.e. the petal primordia are early larger than the stamen primordia, as also in Myrsine and Aegiceras (see especially Ma & Saunders 2003), whereas the stamen primordia may initially be larger, as in Samolus (e.g. Sattler 1962). However, this is a tricky character, since there are really two variables, the relative positions of these priordia and how fast they initially develop, and, as with evicted terminal inflorescences, initial topolgical relationships between parts can speedily become disrupted by post-initiation growth. The number of carpels can be difficult to ascertain, but five seems to be a common number, however, the orientation is unclear. The diagrams presented by Dickson (1936) mostly suggest that the carpels are opposite the petals, but in Primula, at least, they suggest that the carpels are opposite the sepals.

For the hollow style, see Guéguen (1901: is Maesaceae known?), for staminodes, see Saunders (1936) and Caris and Smets (2004: those of Samolus and Theoprastaceae are developmentally rather different), for nectar secretion, see Vogel (1986, 1997) and Caris and Smets (2004), for embryology especially of the herbaceous taxa, i.e. Primulaceae in the old sense, see Dahlgren (1916), for wood anatomy, see Lens et al. (2005a), and for floral morphology and ontogeny, Dickson (1936: esp. gynoecial arrangement), Sattler (1962), Sundberg (1982), Ronse Decraene (1992), Ronse Decraene et al. (1995) and especially Ma and Saunders (2003). For general morphology, see Anderberg et al. (2000) and especially Ståhl and Anderberg (2004).

Phylogeny. The monophyly of the group is not in doubt (see Anderberg & Ståhl 1994; Anderberg et al. 1998; and especially Källersjö et al. 2000: note that support values for Samolus as sister to Theophrastaceae s. str. are reduced when morphological data are added to molecular).

Phylogeny. This whole group was often recognised as Primulales in the past. Perhaps the only question, particularly in light of the break-up of Primulaceae, the removal of Maesa from Myrsinaceae, the placement/addition of Samolus as sister to the old Theophrastaceae, the many herbaceous ex-Primulaceae that are sister to the old-style, woody Myrsinaceae rather than being in a clade with other Primulaceae, and the numerous features shared by the group as a whole, is whether it is worth recognising families at all... Subfamilial names are already available.

Previous relationships. Plumbaginaceae (see Caryophyllales here) were often associated with Primulaceae and related families because of similar placentation and stamen arrangement (see Cronquist 19181 for discussion).

MAESACEAE Anderberg, B. Ståhl & Kallersjö   Back to Ericales

Evergreen lianes or trees; vessel element type?; petiole bundles all annular; secretory canals well developed; leaves spiral or two-ranked, induplicate, margin toothed to entire; inflorescence often branched; flowers small, C induplicate-valvate, stamen primordium smaller than petal primordium, A basally connate, attached at the middle of the C tube, nectary on G; G [3-4], half inferior, ovules apotropous, endothelium +, stigma truncate or capitate and lobed; fruit a many-seeded drupe, K persistent; testa 2-layered, inner layer with rhombic crystals; n = 10.

1/150. Old World tropics to Japan, the Pacific, and Australia (map: from Palgrave 2002).

• Maesaceae are woody plants that can be recognised by their serrate, exstipulate leaves that often lack much venation, even when dry, and which have well-developed and conspicuous canals in the leaf blades, sepals and petals. The small, obviously bracteolate flowers have a connate corolla, stamens opposite the petals and an inferior ovary; the many-seeded fruit is drupaceous.

Chemistry, Morphology, etc. Vessels are in radial multiples (as quite commonly in woody Theophrastaceae and Myrsinaceae); there may be groups of druses in the abaxial epidermis; the fibers are septate; and the lateral bundles arise about half an internode below the leaf they supply. Information on floral development is taken from Caris et al. (2000); the ovules are separated by and partly sunken in placental tissue (see also Utteridge & Saunders 2001).

Theophrastaceae [Myrsinaceae + Primulaceae]: herbs[?]; rays ³5-seriate, uniseriate rays 0 [not herbaceous taxa]; bracteoles 0; C imbricate, arising abaxially on common primordium [i.e. stamen primordium > petal primordium], subrotate, tube rather short.

Phylogeny and Evolution. For the suggestion that rosette herbs may be the plesiomorphic condition for this part of the clade, see Anderberg et al. (2001); however, Lens et al. (2005a) find no evidence from wood anatomy that this is likely (apart from in a few Myrsinaceae). Note that in Myrsinaceae, herbaceous taxa such as Stimpsonia, Ardisiandra and Coris are basal to woody taxa, and variation in habit is very extensive in this clade as a whole. Smith and Donoghue (2008) found that the rate of mollecular evolution in the herbaceous taxa they examined was much greater than in the woody taxa.

THEOPHRASTACEAE Link, nom. cons.   Back to Ericales

Bracts displaced up the pedicels; staminodes +, petaloid, endothelium?

6-9[list]/105 - two groups below. Mostly New World and tropical, some also more temperate and Old World (map: from Hultén 1971).

1. Samolus

Nodes ?1:1; leaves entire; K connate, nectary on ovary; G [5], semi-inferior, style impressed; fruit a 5-valved capsule, seeds many; coat undistinguished, exotesta and endotegmen tanniniferous, the latter crystalliferous; endosperm cell walls thin; n = (12) 13.

1/15. America, the Antipodes, Europe, tropical to temperate (map: from Hultén 1971; Meusel et al. 1978; FloraBase 2005). [Photo - Flowers.]

Chemistry, Morphology, etc. Ståhl (2004) suggests that a secretory system is present, if not always conspicuous. The stomata are anomocytic. There are several petiole bundles forming an arc, and these seem to diverge very soon after the leaf trace departs from the central stele. The ovules completely cover the placenta, but fingers of placental tissue may poke up between them (but not seen in the material examined by Caris & Smets 2004); Ma and Saunders (2003) suggest that in this whole clade (i.e. Theophrastaceae s.l.) the ovules are not embedded (which would then be a synapomorphy for it). The valves of the capsule are opposite the calyx (Caris & Smets 2004).

For general information, see Ståhl (2004: as Samolaceae).

Synonymy: Samolaceae Rafinesque

2. The Rest (Theophrastaceae s. str.)

Woody, tending to be pachycaul; rays broad; nodes also 1:1 [Jacquinia, dividing into three], 5:5 [Clavija]; secretory system?; petiole bundle deeply arcuate or annular, with small adaxial inverted bundles; scale leaves +; leaves conduplicate, margins spiny-toothed to entire, subepidermal fibers +; plant dioecious or flowers bisexual; anthers extrorse, with calcium oxalate, (nectariferous hairs +), style long, stigma dry or wet; fruit a (rather dry) berry, placentae ± pulpy, (drupe); seeds 1-few, rounded, exotestal cells flattened, thick-walled, hypodermal cells (with thickened anticlinal walls), often crystalliferous; endosperm cell walls pitted, cotyledons usu. foliaceous; n = 18, 20, 24.

4/90: Clavija (50), Jacquinia (35 - perhaps to be divided). New World tropics (map: from Ståhl 1989, 1991, 1995). [Photos - Collection]

• Theophrastaceae have sympetalous flowers with stamens opposite the petals and five, petaloid staminodes opposite the sepals. Most taxa are woody plants with broad rays and often pseudoverticillate leaves with toothed or spiny margins or a pungent apex. The extrorse anthers form a cone in the center of the flower when it opens, but later spread; the staminodes are more or less petaloid. The subepidermal fibers are visible in the dried leaf as fine striations - use a hand lens!

Chemistry, Morphology, etc. The subepidermal fibres may lack lignification. For reports of glandular dots on calyx and corolla, see Mabberley (1997). Floral primordia may initially be quite strongly monosymmetric, as in Deherania (Sattler 1962), even if the flower at anthesis is polysymmetric.

For morphology, etc., see Ståhl (2004) and in particular Caris and Smets (2004).

Phylogeny. Phylogenetic relationships suggested by Källersjö and Ståhl (2003) imply that some generic realignments are needed.

Primulaceae + Myrsinaceae: two ndhF deletions.

PRIMULACEAE Borkhausen, nom. cons.   Back to Ericales

Cucurbitacins +; ?cork; glands 0, trichomes articulated; leaves involute or revolute, margins entire to dentate or serrate; inflorescence scapose; K often connate, C hypocrateriform; A attached at or above middle of C tube, pollen syn- or polycolpate, nectary on ovary, ovules not immersed in placenta (immersed - Dionysia), (inner integument ca 4 cells across), style usu. long, (heterostyly +); fruit a capsule; seeds many, angled, exotesta ± persistent, walls thickened or not, (endotesta with inner walls thickened [Primula]), endotegmen often crystalliferous; (endosperm cell walls thin); n = 8-12.

9[list]/900: Primula (490-600: inc. Cortusa, Dionysia [some chasmophytes, for which see Trift et al. 2004, relationships, biogeography; Lidén 2007, revision), Dodecatheon), Androsace (160: inc. Douglasia, Vitaliana, see Schneeweiss et al. 2004b). Northern hemisphere, scattered elsewhere (map: from Hultén 1971; Meusel et al. 1978). [Photo - Dodecatheon flower © R. Kowal] [Photo - Primula flower]

• Primulaceae are usually rosette herbs with a scapose inflorescence and medium-sized flowers with connate sepals, obviously connate petals, hypocrateriform corolla and stamens borne opposite the corolla lobes; the placentation is free-central and the capsular fruits have numerous, angular seeds.

Evolution. Heterostyly is common, although it is unlikely to be an apomorphy for the family; it is sometimes lost, as in those Primula with buzz pollination, the erstwhile Dodecatheon (Mast et al. 2001, 2006).

Chemistry, Morphology, etc. The involute leaves can be sharply bent rather than incurved (for ptyxis, see Conti et al. 2000; Mast et al. 2001). Solereder (1908) reports that secretory tissues occur in Androsace lactea. The corolla epidermal cells are isodiametric. Saunders (1936) suggested that some of the lobing of the corolla of Soldanella might be staminodial.

For pollen variation, see Mast et al. (2001), and for general information, see Anderberg (2004).

Phylogeny. For ITS-based relationships within the family, see Martins et al. (2003), and for relationships within Primula, see also Trift et al. (2002) and Mast et al. (2004, 2006), for relationships within Androsace, see Wang et al. (2004) and Schneeweiss et al. (2004b).

Classification. Richards (2003) provides a good general description of the species of Primula s. str.

MYRSINACEAE R. Brown, nom. cons.   Back to Ericales

Also trees to shrubs or lianes; benzoquinones +; (vessel elements with scalariform perforations); (nodes 3:3 - unnamed taxon from Atlantic Forest; Ardisia densiflora); glands/canals throughout the plant (0); leaves (opposite), also supervolute (curved), margins entire (crenate to serrate, teeth cartilaginous); (plant dioecious), inflorescence often fasciculate/corymbose; flowers (3-)4-5(-7)-merous, C often contorted, (nectariferous hairs +); A dorsifixed or basifixed, sagittate, (porose), (micropyle endostomal - Coris; endothelium 0; style 0; long), stigma (punctate), dry or wet; fruit a berry, drupe or capsule [latter in herbaceous taxa], placentae ± pulpy; seeds 1-few, rounded (ruminate; hilum depressed) [woody taxa] or many, small, angular, seed coat undistinguished, (endotesta crystalliferous - Cyclamen), tegmen thickened before becoming crushed, (endotegmen crystalliferous); endosperm walls pitted, (embryo slightly curved; medium); n = 10-13, 15, 17, 23.

41[list]/1435: Ardisia (450), Myrsine (155: inc. Rapanea, Suttonia, many species in the Pacific), Lysimachia (150: sometimes staminodes?, some woody; white flowers with nectariferous hairs, some yellow oil flowers [Macropis the pollinator], or selfers - see Vogel 1986), Discocalyx (115: inc. Tapeinosperma), Embelia (100), Parathesis (85), Stylogyne (60). Pantropical and N. Temperate (map: from Hultén 1958, 1971; FloraBase 2008: S. Hemisphere a bit notional). [Photos - collection woody members, Cyclamen flower © H. Schneider, fruit © H. Schneider], collection of ex Primulaceae.]

• Myrsinaceae are trees to herbs, and many have distinctive yellowish to blackish dots or streaks on the often spiral leaves and also often obvious on the persistent calyx and on the fruit - I have not seen them on genera like Trientalis, Anagallis (but there may be pigmented spots on the abaxial surface of the lamina), Cyclamen, or some species of Lysimachia. The sympetalous corolla is often contorted, the five stamens are antepetalous, the ovary is superior, and the ovules and/or seeds are immersed in the placenta. In many woody taxa the point of insertion of the branches on the main stem is vertically elongated; growth is often sylleptic.

Evolution. Vogel (1986) discusses pollination, which in a group of yellow-flowered Lysimachia in particular is by oil-collecting Macropis (Mellitidae) bees (see also Simpson et al. 1983). The oil is secreted by trichomes. Anderberg et al. (2007) suggested that Lysimachia with buzz-pollinated flowers and those with nectar-producing hairs formed separate clades and were both derived from oil-producing ancestors.

Some species of Ardisia have pustules along the edge of the leaf blade; although inhabited by bacteria, it is unclear what role the bacteria might be playing (Miller 1990).

For the evolution of the mangrove habitat, to which Aegiceras is restricted, see Rhizophoraceae and Tomlinson (1986). Aegiceras has a number of anomalous anatomical and morphological features, the seed characters in particular are those that might be expected from a mangrove plant, since seeds lack endosperm and contain a large, viviparous embryo (cf. Rhizophoraceae-Rhizophoreae, Acanthaceae-Acantheae-Acanthus ilicifolius, etc.).

Chemistry, Morphology, etc. The presence of coloured glands may well not be a synapomorphy of the family (Hao et al. 2004). There are breakdown areas in the rays of woody members, and these may be filled with dark contents (Lens et al. 2005). Discocalyx has three traces in the petiole base, and some other taxa may be trilacunar; nodal anatomy needs study. The epidermal cells of the corolla are often elongated (as in Glaux - to be included in Lysimachia - lacks a corolla); this is a derived feature within the family. Trientalis has anisomerous flowers (Swenson et al. 2008c). Cyclamen has one cotyledon and one integument. See Oh et al. (2008) for the seed morphology of herbaceous taxa around Lysimachia.

Coris is a particularly distinctive genus morphologically. It is a small ericoid sub-shrub with monosymmetric flowers that have a spine-tipped epicalyx. There is nectary at the base of the ovary and there are only 5-6 ovules. Monosymmetry is expressed early in development by the calyx, but monosymmetry of the corolla becomes evident only later (Ronse Decraene et al. 1995: they suggest that the median sepal is abaxial, i.e. that the orientation of the flower is inverted or oblique). The rays of Aegiceras are relatively narrow, the ovules are unitegmic, etc. (Staåhl å Anderberg 2004; Lens et al. 2005).

Some information is taken from Otegui and Cocucci (1999) and from Ståhl and Anderberg (2004); Lens et al. (2005a) provide much information about wood anatomy.

Phylogeny. Myrsinaceae, previously circumscribed to include only woody taxa, now include Anagallis, Ardisiandra, Asterolinon (?= Lysimachia), Coris, Cyclamen, Glaux, Lysimachia, Pelletiera, Stimpsonia and Trientalis (Anderberg et al. 2000, 2001; the limits of Myrsinaceae are not so clear in Martins et al. 2003, but ITS data alone in the latter study. Anderberg et al. (2007) was particularly interested in the relationships of the herbaceous taxa; the family as a whole had moderate support as being monophyletic (72% jacknife), and Cyclamen, the herbaceous taxa, and the woody taxa then formed a trichotomy. Hao et al. 2004 also provide a phylogeny of the family, although focusing on Lysimachia.

Classification.Generic limits in the woody members in particular are unsatisfactory, but the limits of genera like Lysimachia are also unclear (Anderberg et al. 2007).

Synonymy: Aegicerataceae Blume, Anagallidaceae Adanson, Ardisiaceae Jussieu, Coridaceae J. Agardh, Embeliaceae J. Agardh, Lysimachiaceae Durande