The families of flowering plants.
IncludingLeuchtenbergiaceae Salm-Dyck ex Pfeiff., Nopalaceae (Nopaleaceae) Burnett, Opuntiaceae Martinov
Habit and leaf form. Shrubs, or herbs; non-laticiferous and without coloured juice (usually), or laticiferous (rarely). ‘Normal’ plants (more or less, Pereskieae), or switch-plants, or plants of very peculiar vegetative form (usually more or less spiny, with clusters of spines in hairy, spiralled areoles); ‘cactoid’ (usually), or with the principal photosynthesizing function transferred to stems (sometimes with cylindrical-ridged stems, sometimes with cladodes). Leaveswell developed (Pereskieae), or much reduced to absent (mostly). Plants succulent (mostly), or non-succulent (Pereskieae, with only ‘more or less fleshy’ leaves). Perennial. Self supporting, or epiphytic, or climbing. Pachycaul. Xerophytic. Leaves when present and identifiable as such, deciduous (often caducous); when present, alternate; spiral; ‘herbaceous’, or fleshy, or membranous (usually small and ephemeral); petiolate to sessile; non-sheathing; simple. Lamina entire; one-veined, or pinnately veined; cross-venulate, or without cross-venules.
General anatomy. Plants with laticifers (anastomosing, e.g. in Coryphantha, Leuchtenbergia,Neomamillaria), or without laticifers (usually).
Leaf anatomy. Minor leaf veins without phloem transfer cells (Opuntia, Zygocactus).
Stem anatomy. Secretory cavities present, or absent; when present, with latex, or with mucilage (but mucilage cells much commoner). Cork cambium present; initially superficial. Nodes when recorded, unilacunar (with one trace, this often bi- or multi-fid). Primary vascular tissue comprising a ring of bundles. Cortical bundles present, or absent. Medullary bundles present, or absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent. Xylem without fibre tracheids; with libriform fibres; with vessels (usually), or without vessels. Vessel end-walls simple, or reticulately perforated (rarely). Wood parenchyma typically paratracheal. Sieve-tube plastids P-type; type III (a).
Reproductive type, pollination. Plants hermaphrodite (usually). Pollination anemophilous, or entomophilous, or ornithophilous, or cheiropterophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes. The ultimate inflorescence unit (when flowers aggregated) cymose. Flowers medium-sized to large (often showy); fragrant (often), or odourless; regular to somewhat irregular; when irregular, zygomorphic. The floral irregularity (when noticeable) involving the perianth, or involving the perianth and involving the androecium. Flowerspartially acyclic. The perianth acyclic, or the perianth acyclic and the androecium acyclic. Free hypanthium present, or absent.
Perianthsequentially intergrading from sepals to petals, or petaline; 20–100 (‘many’); free, or joined (basally); green and white, or cream, or yellow, or orange, or red, or pink, or purple.
Androecium 15–100 (‘many’). Androecial members branched, or unbranched; maturing centrifugally; free of the perianth (occasionally), or adnate (to perianth tube); free of one another, or coherent (sometimes in groups). Androecium exclusively of fertile stamens. Stamens 15–100 (‘many’). Anthers dehiscing via longitudinal slits; introrse; bilocular; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with one middle layer. Tapetum glandular. Pollen grains aperturate; 3 aperturate, or 6–15 aperturate; colpate, or foraminate, or rugate; spinulose; 3-celled (recorded in 8 genera).
Gynoecium 3–100 carpelled (to ‘many’). The pistil 1 celled (nearly always), or 2–15 celled (?—Pereskia). Gynoecium syncarpous; synstylovarious; inferior (nearly always), or superior (Pereskia). Ovary 1 locular (occasionally partially partitioned). Locules partially secondarily divided by ‘false septa’ (Pereskia), or without ‘false septa’ (usually). Epigynous disk present (within the hypanthium). Gynoecium stylate. Styles 1; apical. Stigmas 3–100 (as many as G); wet type; non-papillate; Group IV type. Placentation parietal (usually), or basal (Pereskia). Ovules in the single cavity 15–100 (‘many’); long funicled; arillate, or non-arillate; circinotropous, campylotropous (usually), or anatropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (sometimes with filiform apparatus). Endosperm formation nuclear.
Fruit fleshy (usually), or non-fleshy; indehiscent (usually), or dehiscent (rarely); a capsule (rarely), or a berry (usually). Seeds non-endospermic. Perisperm present, or absent. Seeds winged, or wingless. Seeds with starch. Embryo rudimentary at the time of seed release to well differentiated. Cotyledons 2 (when differentiated, free or united). Embryo achlorophyllous (4/5); straight, or curved (usually), or other than straight, curved, bent or coiled (sometimes spiral).
Physiology, biochemistry. Cyanogenic (rarely), or not cyanogenic. Alkaloids present (usually), or absent. Iridoids not detected. Proanthocyanidins absent. Flavonols present, or absent; quercetin, or kaempferol and quercetin. Ellagic acid absent (3 genera). Betalains present. Saponins/sapogenins present, or absent. CAM (nearly always), or C3 (dubiously?). C3 physiology recorded directly in Pereskia, according to Krenzer et al. 1975. CAM recorded directly in Bergerocactus, Carnegiea, Cereus,Cephalocereus, Chamaecereus, Copiapoa, Coryphantha,Echinocactus, Echinocereus, Echinopsis, Eulychnia, Ferocactus,Lobivia, Lophocereus, Machaerocereus, Mammillaria, Melocactus,Myrtillocactus, Neochilena, Neolloydia, Nopalea, Notocactus,Opuntia, Pachycereus, Phyllocactus, Pilocopiapoa, Rhipsalis,Trichocereus, Zygocactus (etc.? — ‘all genera’, according to Smith and Winter, 1996).
Geography, cytology. Temperate, sub-tropical, and tropical. Chiefly in the drier regions of tropical America, but reaching Patagonia and ascending to the High Andes; only Rhipsalis (perhaps introduced) in Africa, Madagascar, Mauritius, Seychelles and Ceylon. X = 11. Supposed basic chromosome number of family: 11.
Taxonomy.Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Caryophylliflorae; Caryophyllales. Cronquist’s Subclass Caryophyllidae; Caryophyllales. APG 3 core angiosperms; core eudicot; Superorder Caryophyllanae; Order Caryophyllales.
Species 2000. Genera about 90 (according to Gunn et al. 1992, other estimates varying from 50 to 150); Acanthocalycium, Acanthocereus, Aporocactus,Ariocarpus, Armatocereus, Arrojadoa, Arthrocereus, Astrophytum,Austrocactus, Aztekium, Bergerocactus, Blossfeldia,Brachycereus, Brasilicereus, Browningia, Calymmanthium,Carnegiea, Cephalocereus, Cereus, Cipocereus, Cleistocactus,Coleocephalocereus, Copiapoa, Corryocactus, Coryphantha,Denmoza, Discocactus, Disocactus, Echinocactus, Echinocereus,Echinopsis, Epiphyllum, Epithelantha, Eriosyce, Escobaria,Espostoa, Espostoopsis, Eulychnia, Facheiroa, Ferocactus,Frailea, Gymnocalycium, Haageocereus, Harrisia, Hatiora,Heliocereus, Hylocereus, Jasminocereus, Leocereus, Lepismium,Leptocereus, Leuchtenbergia, Lophophora, Maihuenia,Mammillaria, Melocactus, Micranthocereus, Mila, Myrtillocactus,Neolloydia, Neoporteria, Neoraimondia, Neowerdomannia,Obregonia, Opuntia, Oreocereus, Pachycereus, Parodia,Pediocactus, Pelecyphora, Peniocereus, Pereskia, Pereskiopsis,Pilosocereus, Pseudorhipsalis, Pterocactus, Rebutia,Rhipsalis, Samaipaticereus, Schlumbergera, Sclerocactus,Selenicereus, Stenocactus, Stenocereus, Stephanocereus,Stetsonia, Strombocactus, Tacinga, Thelocactus, Uebelmannia,Weberbauerocereus, Weberocereus, Zygocactus.
Economic uses, etc. Some edible fruits (prickly pear, India fig, tuna, arridari, pitaya).