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). ‘Normal’ plants. Plants non-succulent (usually), or succulent. Annual, or biennial. 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. Stipules not ochreate.
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. Plants 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 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.
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, biochemistry. Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. 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 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).
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 (1998) Eudicot; core Eudicot; neither Rosid nor Asterid; 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. • Silenoideae (Agrostemma, Silene). • Silenoideae (Silene). • Silenoideae (Dianthus), Alsinoideae (Scleranthus). • Alsinoideae (Myosoton, Cerastium). • Alsinoideae (Stellaria, Moehringia). • Alsinoideae (Sagina, Honkenya, Moenchia, Arenaria, Spergularia). • Technical details (Silene, Cucubalus, Viscaria, Agrostemma). • Technical details (Dianthus, Velezia, Gypsophila). • Technical details (Stellaria, Cerastium, Colobanthus,Holosteum). • Technical details (Polycarpaea). • Technical details (Scleranthus, Paronychia). • Technical details (Illecebrum, Dicheranthus). • `Franklin's Tartar' (virus infected carnation).
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’)
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).
