Plumbaginaceae Juss.
Including Aegialitidaceae Lincz., Armeriaceae Horan., Limoniaceae Lincz., Pentaptychaceae Dulac, Staticinae (Staticaceae) Hoffmgg. & Link ex S.F. Gray
Habit and leaf form. Herbs (usually), or shrubs, or lianas. Perennial; with a basal aggregation of leaves, or with neither basal nor terminal 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; without a persistent basal meristem.
Leaf anatomy. Stomata anomocytic, or anisocytic, or paracytic.
The mesophyll with sclerencymatous idioblasts. Minor leaf veins with phloem transfer cells (7 genera).
Stem anatomy. Cork cambium present; initially superficial. Nodes tri-lacunar. Cortical bundles often present. Medullary bundles often present. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous (often); commonly via concentric cambia. ‘Included’ phloem commonly present. Xylem with libriform fibres. Vessel end-walls simple. Wood parenchyma scanty paratracheal. Sieve-tube plastids S-type.
Reproductive type, pollination. Plants hermaphrodite; heterostylous (often, and the pollen often dimorphic in Armerioideae), or heterostylous (?). 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 terminal inflorescence unit 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 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, biochemistry. Cyanogenic (Plumbago), or not cyanogenic (mostly). Alkaloids present, or absent. Iridoids not detected. 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). Arbutin absent. Betalains absent. Saponins/sapogenins absent. Aluminium accumulation not found. C3. C3 physiology recorded directly in Limoniastrum, Limonium. Anatomy non-C4 type (Acantholimon, Limonium, Psilliostachys).
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 (1998) Eudicot; core Eudicot; neither Rosid nor Asterid; Caryophyllales. Species 775. Genera 24; Acantholimon, Aegialitis, Armeria, Bamiana, Buciniczea, Cephalorhizum, Ceratostigma, Chaetolimon, Dictyolimon, Dyerophytum, Eremolimon, Ghasnianthus, Goniolimon, Ikonnikovia, Limoniastrum, Limoniopsis, Limonium, Meullerolimon, Neogontscharovia, Plumbagella, Plumbago, Popoviolimon, Psylliostachys, Vasssilczenkoa.
Illustrations. • Limonium vulgare. • Armeria maritima. • Limonium sinuatum. • Technical details (Plumbago, Statice, Armeria). • Technical details (Dyerophytum). • Technical details (Armeria).
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)
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
