| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Plumbaginaceae Juss.

Including Aegialitidaceae Lincz., Armeriaceae Horan., Limoniaceae Lincz., Pentaptychaceae Dulac, Staticinae (Staticaceae) Hoffmgg. & Link ex S.F. Gray.

Habit and leaf form. Divers in habit - herbs (mostly), or trees (small), or ‘arborescent’, or shrubs, or lianas. Perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves. Self supporting, or climbing; sometimes stem twiners; Plumbago twining clockwise. Helophytic, or mesophytic, or xerophytic (especially salt steppes and maritime). Leaves alternate; spiral; ‘herbaceous’, or leathery (Aegialitis); petiolate to sessile; sheathing (Eagialitis), or non-sheathing; simple; epulvinate. Lamina dissected, or entire; often acicular, or linear, or oblong (narrow); when dissected, pinnatifid; pinnately veined, or palmately veined to parallel-veined (Aegialitis). Leaves stipulate, or exstipulate; leaf development not ‘graminaceous’.

Leaf anatomy. The leaf lamina variously dorsiventral, or bifacial, or centric (characteristically bearing conspicuous epidermal glands secreting mucilage and/or calcium salts). Stomata present; mainly confined to one surface, or on both surfaces; anomocytic, or anisocytic, or paracytic (in narrow-leaved species). Hairs present; eglandular (mostly, but not exclusively, unicellular and simple), or eglandular and glandular (sometimes with long-stalked, shaggy glandular hairs as well, these additional to the peculiar epidermal glands). The mesophyll with sclerenchymatous idioblasts; containing crystals (but these very rare), or without crystals. The crystals when recorded, druses, or solitary-prismatic. Minor leaf veins with phloem transfer cells (found in 7 genera).

Axial (stem, wood) anatomy. Secretory cavities absent (but secretory cells with assumed tanniniferous contents or plumbaginin common). Cork cambium present; initially superficial. Nodes tri-lacunar. Primary vascular tissues comprising a ring of bundles; collateral. Internal phloem absent. Cortical bundles present (often), or absent. Medullary bundles present (sometimes inversely orientated or centric), or absent. Secondary thickening developing from a conventional cambial ring, or anomalous (often). The anomalous secondary thickening when present, via concentric cambia, or from a single cambial ring.

The wood diffuse porous. The vessels very small; solitary, radially paired, and in radial multiples (the multiples short). The vessel end-walls simple. The axial xylem without fibre tracheids; with libriform fibres; including septate fibres (rarely), or without septate fibres. The parenchyma scanty paratracheal. ‘Included’ phloem of the concentric type present, or absent. The wood not storied.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite; heterostylous (often, and the pollen often dimorphic in Armerioideae), or heterostylous (?). Pollination entomophilous; via hymenoptera, via lepidoptera, and via diptera.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in panicles, or in heads, or in racemes. The ultimate inflorescence units cymose, or racemose. Inflorescences scapiflorous, or not scapiflorous; variously panicles or cymose heads (Armerioideae) or racemes (Plumbaginoideae). Flowers bracteolate; small; regular; 5 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk absent.

Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous; sepaloid and petaloid, or sepaloid (the calyx often petaloid). Calyx 5; 1 whorled; gamosepalous (forming a 5 or ten ribbed tube); blunt-lobed, or toothed. Calyx lobes markedly shorter than the tube. Calyx regular; persistent; valvate, or plicate in bud. Corolla 5; 1 whorled; gamopetalous, or polypetalous (or almost so); imbricate, or contorted; regular; white, or yellow, or red (or rose), or pink, or purple, or blue, or red and blue; fleshy (or leathery, in Aegialitis), or not fleshy; persistent (often), or deciduous (deciduous above and persistent below in Aegialitis). Petals when free clawed, or sessile.

Androecium 5. Androecial members free of the perianth, or adnate (to the corolla, in Armerioideae); free of one another, or coherent (forming a persistent tube in Aegialitis); 1 whorled. Androecium exclusively of fertile stamens. Stamens 5; isomerous with the perianth; alternisepalous; opposite the corolla members. Anthers dorsifixed, or basifixed (Aegialitis); versatile, or non-versatile; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer. Tapetum glandular (?). Pollen grains aperturate; 3–5 aperturate, or 4 aperturate, or 6 aperturate; colpate (3–5, or 4-rupate), or rugate (6-, or more or less irregular); 3-celled.

Gynoecium 5 carpelled. Carpels isomerous with the perianth. The pistil 1 celled. Gynoecium syncarpous; synovarious (Armerioideae, Aegialitis), or synstylovarious to eu-syncarpous; superior. Ovary 1 locular. Styles 1, or 5 (opposite the sepals); when five, free to partially joined; apical. Stigmas 5; dry type; papillate; Group II type. Placentation basal. Ovules in the single cavity 1; pendulous (from a long, basally attached funicle); anatropous (or circinnotropous); bitegmic; crassinucellate. Embryo-sac development Adoxa-type, or Penaea-type, or Fritillaria-type, or Plumbago-type. Endosperm formation nuclear. Embryogeny solanad.

Fruit dehiscent, or indehiscent; a capsule, or a nut. Capsules when dehiscent circumscissile, or valvular (with apical valves). Seeds endospermic, or non-endospermic; winged. Embryo well differentiated. Cotyledons 2; flat. Embryo chlorophyllous (4/5); straight.

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Limoniastrum, Limonium. Anatomy non-C4 type (Acantholimon, Limonium, Psilliostachys). Cyanogenic (Plumbago), or not cyanogenic (mostly). Alkaloids present, or absent. Arbutin absent. Iridoids not detected. Betalains absent (where sought). Saponins/sapogenins absent. Proanthocyanidins present; delphinidin, or cyanidin and delphinidin. Flavonols present; kaempferol and myricetin, or quercetin and myricetin, or kaempferol, quercetin, and myricetin. Ellagic acid absent (3 genera, 6 species). Aluminium accumulation not found. Sieve-tube plastids S-type.

Geography, cytology. Frigid zone to tropical. Cosmopolitan, especially salt steppes and coasts. X = 6–9 (mostly).

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Malviflorae; Plumbaginales. Cronquist’s Subclass Caryophyllidae; Plumbaginales. APG III core angiosperms; core eudicot; Superorder Caryophyllanae. APG IV Order Caryophyllales.

Species 775. Genera about 25; Acantholimon, Aegialitis, Afrolimon, Armeria, Bamiana, Buciniczea, Cephalorhizum, Ceratostigma, Chaetolimon, Dictyolimon, Dyerophytum, Eremolimon, Ghasnianthus, Goniolimon, Ikonnikovia, Limoniastrum, Limoniopsis, Limonium, Meullerolimon, Neogontscharovia, Plumbagella, Plumbago, Popoviolimon, Psylliostachys, Vasssilczenkoa.

Quotations.

The marsh is bleak and lonely. Scarce a flower
Gleams in the waving grass. The rosy thrift
Has paler grown since summer bless’d the scene,
And the Sea Lavender, whose lilac blooms
Drew from the soil a richer hue
Than when they grew on yonder towering cliff,
Quivers in flowerless greenless to the wind
(Quoted by Ann Pratt, ‘Wild Flowers’ (1857), unattributed - Armeria, Limonium)

Illustrations. • Le Maout and Decaisne: Plumbago, Statice, Armeria. • Acantholimon venustum: Bot. Mag. 122 (1896). • Acantholimon ecae: Trans. Lin. Soc. Lond. Bot. 3 (1887). • Armeria caespitosa: Bot. Mag. 124 (1898). • Armeria maritima (as A. vulgaris): Lindley. • Armeria maritima (as A. vulgaris) and A. arenaria (as A. plantaginea): Eng. Bot. 1152–1154, 1867. • Armeria maritima (B. Ent.). • Armeria mauritanica (as A. cephalotes): Bot. Mag. 70 (1844). • Armeria pungens: as A. fasciculata, Bot. Reg. 21, 1841. • Armeria pseudoarmeria (as A. latifolia): Bot. Mag. 119 (1893). • Ceratostigma plumbaginoides, as Valoradia: Bot. Mag. 76 (1850). • Ceratostigma willmottianum: Bot. Mag. 140 (1914). • Dyerophytum africanum: Thonner. • Goniolimon tataricum (as Statice): Bot. Mag. 107 (1881). • Limoniastrum articulatum: as Statice monopetala, Bot. Reg. 54, 1841. • Limonium arboreum (as Statice arborea): Bot. Mag. 66 (1840). • Limonium bellidifolium (as Statice caspia): Eng. Bot. 11611, 1867. • Limonium bonduellei (as Statice): Bot. Mag. 86 (1869). • Limonium brassicifolium (as Statice): Bot. Mag. 86 (1860). • Limonium humile (as Statice bahusiensis) and two forms of L. binervosa (as Statice binervosa): Eng Bot. 1158, 1867. • Limonium macrophyllum (as Statice macrophylla): Bot. Mag. 70 (1844). • Limonium pectinatum: as Statice, Bot. Reg. xxvi, 65 (1840). • Limonium puberulum: as Statice puberula, Bot. Reg. 1450, 1831. • Limonium rytidophylla (as Statice): Bot. Mag. 70 (1843). • Limonium sinuatum, as Statice: Bot. Mag. 2 (1788). • Limonium vulgare (B. Ent.). • Plumbago indica (as P. rosea var. coccinea): Bot. Mag. 89 (1863). • Plumbago zeylanica: Wight, Illustrations of Indian Botany 2 (1850). • Psylliostachys suvorovii, as Statice: Bot. Mag. 113 (1887). • Limonium puberulum (as Statice): Bot. Mag. 65 (1839). • Glandular leaf hairs of Aegialitis annulata: Solereder, 1908.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 8th June 2026. delta-intkey.com’.

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