Plumbaginaceae  

Perennial or sometimes annual herbs, subshrubs, or low shrubs, rarely lianas. Secondary growth in some genera (including Limoniastrum) anomalous. Vessels with simple perforations; lateral pitting alternate. Fibers with simple pits. Rays homogeneous. Axial parenchyma scanty paratracheal. Sieve-element plastids of S-type. Leaves alternate, entire, usually estipulate, only rarely with well-developed stipules, often with scattered chalk-glands exuding water and salts, and often also raised mucilage glands in leaf axils and on the surface. Stomata anomocytic, or anisocytic, or paracytic. Flowers in cymose heads or panicles or in racemes or spikes, bisexual, actinomorphic, 5-merous, with both sepals and petals, bracteate (the bracts sometimes forming on involucre), with 2(1) bracteoles, often heterostylic. Sepals connate to form a plicate, often conspicuously 5-ribbed or 10-ribbed tube, the lobes mostly dry and membranous or scarious, often showy, persistent. Petals connate at the very base or less often into a tube, often persistent; the petals or corolla lobes contorted or imbricate. Stamens five, opposite the petals or corolla lobes, or partly adnate to the tube; anthers dorsifixed or (Aegialitis) basifixed, tetrasporangiate, introrse, 2-locular, opening longitudinally. Tapetum secretory.

Pollen grains 2-celled or more often 3-celled, tectate-columellate, 3-colpate or rarely 4–5-colpate, very rarely pantocolpate, usually dimorphic in the Staticoideae. Gynoecium of five united carpels; stylodia more or less distinct (Staticoideae) or connate into an apically lobed style (Plumbaginoideae); stigmas papillate; ovary superior, unilocular, usually 5-lobed or -ribbed, with solitary basal ovule on elongate, slender funicle that almost encircles it. Ovules pendulous, anatropous, sometimes circinotropous, bitegmic, crassinucellate. Endosperm nuclear. Fruits mostly more or less enclosed within the persistent calyx, dry, indehiscent or less often tardily circumscissally dehiscent or splitting upward into valves. Seeds small, winged; seed coat formed by both integuments and consists of exotesta and endotegmen (Vovk 1991). Embryo rather large, green, straight, well differentiated; endosperm starchy, more or less copious or scanty.  Plants often salt tolerant; producing various 0-methylated flavonols and anthocyanidins, glycine betaine, choline-O-sulphate, plumbagin, kaempferol, quercetin, and myricetin, but no betalains, generally tanniniferous. n = 6–9. Friedrich (1956) brought together many gross morphological, anatomical, palynological, embryological, and cytological data that confirm the affinity of the Plumbaginaceae with the families of Centrospermae.

PLUMBAGINOIDEAE
Flowers in racemes or spikes. Sepals herbaceous, more or less glandular. Corolla sympetalous. Stamens mostly free or partly connate to the corolla. Pollen grains with well-defined foot-layer, highly irregular columellae, and a complete tectum. Stylodia connate into an apically lobed style. Naphtoquinone plumbagin, glycine betaines, 5-O-methylated flavonols always present, n = 6–7. –Plumbago, Plumbagella.

STATICOIDEAE
Flowers in panicles or heads with monochasial partial inflorescences or rarely (Aegialltis) in leafy, often fork branched racemes. Sepals scarious. Petals connate only at the very base, rarely (Psylliostachys) corolla sympetalous. Filaments basally adnate to the petals. Pollen grains with a foot-layer supporting straight, regular columellae, which are distally fused into an incomplete rectum. Stylodia free or rarely (Limoniastrum) connate up to the middle. Present betaalanine betaines, glycine betaines rare, naphtoquinone plumbagin lacking, n = 8, 9. –Limoniastrum (including Bubania), Limonium.