| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Caryophyllaceae Juss.

Including Alsinaceae, Circumaceae Dulac, Corrigiolaceae Dum., Dianthaceae von Vest, Gracilicaulaceae Dulac, Illecebraceae R. Br., Onychiaceae Dulac, Paronychieae (Paronychiaceae) A. St.-Hil., Scleranthaceae Bartl., Sileneae (Silenaceae) Bartl., Stellariaceae Dum.

Habit and leaf form. Small trees (a few), or shrubs (a few), or herbs (mainly). Plants non-succulent (usually), or succulent; autotrophic (mostly, although numerous species have sticky hairs that trap insects), or carnivorous (? - possibly, since some Cerastium, Silene, Stellaria species are claimed to secrete proteases); green and photosynthesizing. Trapping mechanism if operative, passive. The traps if applicable, consisting of the sticky-glandular, non-irritable (flypaper-like) leaves, or consisting of the sticky-glandular, non-irritable (flypaper-like) inflorescence branches or pedicels. Annual, or biennial, or perennial. Helophytic to xerophytic. Leaves opposite (usually), or whorled (rarely), or alternate (rarely spiral); petiolate, or subsessile, or sessile, or perfoliate; connate, or not connate; simple; epulvinate. Lamina entire (entire). Leaves stipulate (often), or exstipulate.

General anatomy. Plants with ‘crystal sand’ (e.g., in Gymnocarpos), or without ‘crystal sand’.

Leaf anatomy. The leaf lamina dorsiventral (with abaxial palisade in some Arctic species), or centric. Stomata present; mainly confined to one surface, or on both surfaces; diacytic (mostly), or anisocytic and diacytic. Hairs present; eglandular, or eglandular and glandular; unicellular, or multicellular (mostly simple, uniseriate or unicellular, but branched hairs are common in some genera). The mesophyll usually containing crystals. The crystals commonly druses. Minor leaf veins without phloem transfer cells (10 genera).

Axial (stem, wood) anatomy. Cork cambium present, or absent; initially deep-seated (usually), or initially superficial (e.g., in Paronychia). Nodes unilacunar. Primary vascular tissues variously in a cylinder, without separate bundles, or comprising a ring of bundles, or comprising two or more rings of bundles. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous. The anomalous secondary thickening 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. Primary medullary rays often wide.

The vessel end-walls simple. The axial xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres. The parenchyma often constituting much of the wood. ‘Included’ phloem present, or absent. The wood not storied.

Reproductive type, pollination. Unisexual flowers present, or absent. Plants hermaphrodite, or andromonoecious, or dioecious, or gynodioecious. Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’ (usually), or solitary; when aggregated, in cymes. The ultimate inflorescence units 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 median 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, or markedly unequal; 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; spinulose; 3-celled (10 genera).

Gynoecium 2–5 carpelled. The pistil 1–5 celled. Gynoecium 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). The radicle dorsal (nearly always), or lateral (very rarely).

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. C3, or C4. C3 physiology recorded directly in Arenaria, Cerastium, Dianthus, Gymnocarpos, Lychnis, Polycarpon, Silene, Spergularia, Stellaria, Tunica. C4 physiology recorded directly 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). Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. Betalains absent (where sought). Saponins/sapogenins present (commonly), or 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). Sieve-tube plastids P-type; type III (b).

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

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Caryophylliflorae; Caryophyllales. Cronquist’s Subclass Caryophyllidae; Caryophyllales. APG III core angiosperms; core eudicot; Superorder Caryophyllanae. APG IV Order 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, Viscaria, 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).

Quotations.

Quaff’d off the muscadel,
And threw the sops all in the sexton’s face
(‘Taming of the Shrew’, iii., 2 - ‘sops’ (‘sops-in-wine’) = carnations, used for flavouring the sweet wine presented to brides after medieval wedding ceremonies)

Ragged-robbins once so pink
Now are turned as black as ink,
And their leaves being scorch’d so much
Even crumble to the touch
(John Clare 1820, ‘Noon’)

Illustrations. • Le Maout and Decaisne: Agrostemma, Cucubalus, Drypis, Melandrium (= Silene), Viscaria. • Le Maout and Decaisne: Gypsophila, Saponaria, Stellaria. • Le Maout and Decaisne: Bufonia, Dianthus, Cerastium, Colobanthus, Holosteum, Spergularia, Stellaria. • Le Maout and Decaisne: Scleranthus, Anychia (= Paronychia). • Le Maout and Decaisne: Illecebrum, Dicheranthus. • Agrostemma githago: Köhler’s Medizinal-Pflanzen 4 (1898). • Agrostemma githago (as Lychnis githago): Eng. Bot. 215, 1864. • Arenaria littledalei (as Gooringia): Hook. Ic. Pl. 30 (1911). • Arenaria norvegica: Eng. Bot. 237, 1864. • Arenaria purpurascens: Bot. Mag. 96 (1870). • Cucubalus baccifer (as C. bacciferus): Eng. Bot. 198, 1864. • Cerastium diffusum (as C. tetrandrum), C. glomeratum and C. arcticum (as C. latifolium): Eng. Bot. 218, 221 and 224, 1864. • Cerastium alpinum, C. arvense and C. cerastoides (as C. trigynum): Eng. Bot. 223, 225 and 226, 1864. • Dianthus armeria, D. deltoides, D. gratianopolitanus (as D. caesius), D. caryophyllus: Eng. Bot. 191–194, 1864. • Dianthus chinensis var. laciniatus: Bot. Mag. 91 (1865). • Gypsophila cerastioides: Bot. Mag. 109 (1883). • Holosteum umbellatum: Eng. Bot. 216, 1864. • Honckenya peploides (as Alsine peploides): Eng. Bot. 239, 1864. • Lychnis bungeana: Bot. Mag. 64 (1837). • Lychnis flos-cuculi and L. viscaria: Eng. Bot. 212–213, 1864. • Lychnis coronata: as L. bungeana, Bot. Reg. 1864 (1836). • Minuartia rubella (as Alsine rubella), M. stricta (as Alsine uliginosa), M. hybrida ssp. tenuifolia (as Alsine tenuifolia): Eng. Bot. 242–244, 1864. • Minuartia rubra ssp. fastigiata (as Alsine fastigiata): Eng. Bot. 243, 1864. • Minuartia sedoides (= Cherleria sedoides, as Alsine cherleria): Eng. Bot. 240, 1864. • Moenchia erecta (as Cerastuim quaternellum): Eng. Bot. 217, 1864. • Moehringia trinervia (as Arenaria trinervia): Eng. Bot. 234, 1864. • Myosoton aquaticum (as Stellaria aquatica): Eng. Bot. 227, 1864. • Paronychia illecebroides: Hook. Ic. Pl. 8 (1848). • Polycarpaea linearifolia: Thonner. • Polycarpon tetraphyllum: Eng. Bot. 258, 1864. • Sagina subulata, S. procumbens and S. nodosa: Eng. Bot. 248–251, 1864. • Saponaria officinalis: Köhler’s Medizinal-Pflanzen 4 (1898). • Saponaria officinalis: Eng. Bot. 197, 1864. • Silene acaulis: Eng. Bot. 205, 1864. • Silene elisabethae: Bot. Mag. 89 (1863). • Silene fissipetala (as S. fortunei): Bot. Mag. 125 (1899). • Silene hookeri: Bot. Mag. 99 (1873). • Silene vulgaris (as S. inflata), S. maritima, S. conica: Eng. Bot. 199–201, 1864. • Silene gallica (as S. anglica and S. quinquevulnera): Eng. Bot. 202–203, 1864. • Silene glaucifolia (as Lychnis lagascae): Bot. Mag. 94 (1868). • Silene nutans and S. noctiflora: Eng. Bot. 207 and 209, 1864. • Silene latifolia (as S. pratensis) and S. dioica (as S. diurna): Eng. Bot. 210–211, 1864. • Silene coeli-rosa: as Viscaria oculata, Bot. Reg. 29, 53 (1843). • Spergula arvensis: Eng. Bot. 253, 1864. • Spergularia rubra and S. media (as S. marginata): Eng. Bot. 254 and 257, 1864. • Sphaerocoma hookeri, as Psyllothamnus beevori: Hook Ic. Pl. 15 (1885). • Stellaria nemorum, S. media and S. uliginosa: Eng. Bot. 228, 229 and 233, 1864. • Stellaria holostea and Stellaria graminea: Eng. Bot. 230 and 232, 1864. • Viscaria oculata: Bot. Mag. 70 (1844). • Xerotia arabica: Hook. Ic. Pl. 24 (1895). • British Silenoideae (Agrostemma, Silene: B. Ent. compilation, 1824–35). • British Silenoideae (Silene: B. Ent.). • British Silenoideae (Dianthus), Alsinoideae (Scleranthus): B. Ent. compilation, 1824–35. • British Alsinoideae (Myosoton, Cerastium: B. Ent, compilation, 1824–35). • British Alsinoideae (Stellaria, Moehringia: B. Ent. compilation, 1824–35). • British Alsinoideae (Sagina, Honkenya, Moenchia, Arenaria, Spergularia: B. Ent. compilation, 1824–35). • ‘Franklin's Tartar’ (freak cultivated Carnation: Bot. Mag. 2 (1788).

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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