Caryophyllaceae Juss.

Including Circumaceae Dulac, Corrigiolaceae Dum., Gracilicaulaceae Dulac, Illecebraceae R. Br., Onychiaceae Dulac, Paronychi(ac)eae A. St.-Hil., Scleranthaceae Bartl., Silen(ac)eae Bartl., Silen(ac)eae Bartl., Stellariaceae Dum.

Habit and leaf form. Small trees (a few), or shrubs (a few), or herbs (mainly). Plants non-succulent (usually), or succulent. Annual, or biennial. Helophytic to xerophytic. Leaves opposite (usually), or whorled (rarely), or alternate (rarely spiral); sessile, or perfoliate; connate, or not connate; simple; epulvinate. Lamina entire (entire). Leaves stipulate (often), or exstipulate.

Leaf anatomy. Minor leaf veins without phloem transfer cells (10 genera).

Stem anatomy. Cork cambium present, or absent; initially deep-seated. Nodes unilacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous; sometimes via concentric cambia (e.g. Spergularia, and sometimes (e.g. Acanthophyllum) producing a second series of bundles internally), or from a single cambial ring. `Included' phloem present, or absent. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls simple. Sieve-tube plastids P-type; type III (b).

Reproductive type, pollination. Hermaphrodite, or andromonoecious, or dioecious, or gynodioecious. Entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in `inflorescences' (usually), or solitary; in cymes. The terminal inflorescence unit cymose. Inflorescences usually terminal; typically dichasial cymes going over into cincinni. Flowers regular; generally more or less 5 merous; cyclic; tetracyclic, or pentacyclic. Floral receptacle developing a gynophore (often), or with neither androphore nor gynophore. Hypogynous disk present.

Perianth with distinct calyx and corolla, or sepaline (occasionally apetalous); (8-)10, or 5; 2 whorled (usually), or 1 whorled; isomerous. Calyx (4-)5; 1 whorled; polysepalous, or gamosepalous; regular; imbricate; with the odd member posterior. Corolla when present, (4-)5; 1 whorled; appendiculate (often, with an appendiculate scale above each petal claw), or not appendiculate; polypetalous; regular. Petals clawed (often), or sessile; deeply bifid to bilobed (often), or fringed, or entire.

Androecium (1-)5, or 8, or 10. Androecial members free of the perianth, or adnate (sometimes adnate to the base of the corolla or the calyx); all equal; free of one another; 1 - whorled, or 2 - whorled. Androecium exclusively of fertile stamens. Stamens (1-)5, or 8, or 10; reduced in number relative to the adjacent perianth (rarely), or isomerous with the perianth, or diplostemonous; oppositisepalous (usually), or alternisepalous (Colobanthus). Anthers dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or decussate. Anther wall initially with one middle layer; of the `dicot' type, or of the `monocot' type (?). Tapetum glandular. Pollen grains aperturate; 3-12 - aperturate; colpate (and sometimes rupate), or porate, or foraminate, or rugate; 3-celled.

Gynoecium 2-5; syncarpous; synovarious to eu-syncarpous; superior. Ovary 1 locular (at least distally, but often more or less partitioned below or when immature). Styles 2-5; free to partially joined; apical. Stigmas 2-5; dry type; papillate; Group II type. Placentation basal, or free central (generally, but the placenta sometimes reaching the apex). Ovules in the single cavity (1-)100 (i.e. occasionally few, usually `many'); ascending; non-arillate; hemianatropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed, or not formed (then the three nuclei degenerating early); when formed, 3; not proliferating; ephemeral. Synergids pear-shaped, or hooked (e.g. Spergula). Endosperm formation nuclear. Embryogeny caryophyllad, or solanad.

Fruit non-fleshy; dehiscent, or indehiscent; a capsule (usually), or a nut (occasionally). Capsules usually denticidal, or valvular. Seeds endospermic, or non-endospermic. Endosperm not oily. Perisperm present. Seeds with starch. Cotyledons 2. Embryo achlorophyllous (14/18); curved (usually), or coiled, or straight (or almost so).

Seedling. Germination phanerocotylar.

Physiology, biochemistry. Not cyanogenic. Alkaloids present, or absent. Iridoids absent. Proanthocyanidins nearly always absent (or a trace of cyanidin in Arenaria). Flavonols present, or absent; when present, kaempferol, or kaempferol and quercetin. Ellagic acid absent (14 species, 11 genera). Betalains absent. Saponins/sapogenins present (commonly), or absent. C3 and C4. C3 recorded in Arenaria, Cerastium, Dianthus, Gymnocarpos, Lychnis, Polycarpon, Silene, Spergularia, Stellaria, Tunica. C4 recorded in Polycarpaea. Anatomy non-C4 type (Acanthophyllum, Arenaria, Cerastium, Dianthus, Drymaria, Gymnocarpos, Lychnis, Polycarpon, Pollichia, Sagina, Silene, Spergula, Spegularia, Stellaria, Uebelina), or C4 type (Polycarpaea).

Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = 5-19.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren's Superorder Caryophylliflorae; Caryophyllales. Cronquist's Subclass Caryophyllidae; Caryophyllales. Takhtajan's Subclass Caryophyllidae; Caryophyllanae; Caryophyllales. Species 1750. Genera 88; Acanthophyllum, Achyronychia, Agrostemma, Allochrusa, Alsinidendron, Ankyropetalum, Arenaria, Bolanthus, Bolbosaponaria, Brachystemma, Bufonia, Cardionema, Cerastium, Cerdia, Colobanthus, Cometes, Cucubalus, Cyathophylla, Dianthus, Diaphanoptera, Dicheranthus, Drymaria, Drypis, Geocarpon, Gymnocarpos, Gypsophilla, Habrosia, Haya, Herniaria, Holosteum, Honckenya, Illecebrum, Kabulia, Krauseola, Kuhitangia, Lepyrodiclis, Lochia, Loeflingia, Lychnis, Mesostemma, Microphyes, Minuartia, Moehringia, Moenchia, Myosoton, Ochotonophila, Ortegia, Paronychia, Pentastemonodiscus, Petrocoptis, Petrorhagia, Philippiella, Phrynella, Pinosia, Pirinia, Pleioneura, Plettkia, Pollichia, Polycarpaea, Polycarpon, Polytepalum, Pseudostellaria, Pteranthus, Pycnophyllopsis, Pycnophyllum, Reicheella, Sagina, Sanctambrosia, Saponaria, Schiedea, Scleranthopsis, Scleranthus, Sclerocephalus, Scopulophila, Selleola, Silene, Spergula, Spergularia, Sphaerocoma, Stellaria, Stipulicida, Thurya, Thylacospermum, Uebelinia, Vaccaria, Velezia, Wilhelmsia, Xerotia.

Economic uses, etc. Numerous ornamentals, e.g. 70 or more species of Dianthus (including carnation), Gypsophila, Silene, Lychnis, and some widespread weeds (Cerastium, Arenaria).

Illustrations. caryo255.gif caryo257.gif caryo258.gif caryo196.gif caryo642.gif caryo643.gif