The World Flora Online

SELECTED REFERENCES

Anderson, E. F. 2001. The Cactus Family. Portland. Barthlott, W. and D. R. Hunt. 1993. Cactaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 4+ vols. Berlin etc. Vol. 2, pp. 161-197. Benson, L. D. 1969b. Cactaceae. In: C. L. Lundell. 1942-1969. Flora of Texas. 3 vols. in parts. Dallas and Renner, Tex. vol. 2, pp. 221-317. Benson, L. D. 1982. Cacti of the United States and Canada. Stanford. Gibson, A. C. and P. S. Nobel. 1986. The Cactus Primer. Cambridge, Massachusetts. Hunt, D. R., comp. 1999. CITES Cactaceae Checklist, ed. 2. Kew. Hunt, D. R. and N. P. Taylor, comps. 1990. The genera of Cactaceae: Progress towards consensus. Bradleya 8: 85-107. Wallace, R. S. 1995. Molecular systematic study of the Cactaceae: Using chloroplast DNA variation to elucidate cactus phylogeny. Bradleya 13: 1-12. Wallace, R. S. and A. C. Gibson. 2002. Evolution and systematics. In: P. S. Nobel, ed. 2002. Cacti: Biology and Uses. Berkeley. Pp. 1-21.

Trees, shrubs, subshrubs, or somewhat vinelike, solitary to forming mats or clumps, terrestrial (sometimes deep-seated in substrate) to epiphytic or epipetric, erect to sprawling (rarely scrambling or climbing) or pendent in epiphytic or epipetric taxa, simple to many branched, usually stem succulent. Roots diffuse, taproots, or tuberlike, sometimes adventitious. Stems unsegmented or segmented, segments persistent to easily detachable; long shoots spheric to depressed-spheric or club-shaped to long cylindric, or sometimes flattened cladodes, smooth, tuberculate and/or fluted with ribs; tubercles distinct as nipple-shaped or ridgelike (to triangular or pyramidal) protuberances to coalescent as vertical ribs; ribs 2-30[-40+], if ribs 2, stems winged, if ribs 3 or more, stems ± angled; short shoots (areoles) positioned on crests of ribs, at or near tubercle apices, or in axils of tubercles, commonly bearing persistent spines, also minute, barbed, deciduous spines (glochids) in subfam. Opuntioideae, and abundant, dense hairs (wool) creating a cushionlike appearance. Leaves deciduous to persistent, vestigial or absent, spirally alternate, sessile (petiolate to subsessile in Pereskia and several genera outside the flora), terete or flat, 0-3 cm (to 10 cm in Pereskia); stipules absent. Spines flexible and hairlike or bristlelike to rigid and needlelike or nail-like, terete to angled or flat, mostly hard (rarely corky or papery). Flowers bisexual (rarely unisexual or with bisexual and pistillate flowers on separate plants), nocturnal or diurnal, 1(-several) per areole, arranged in true inflorescence only in subfam. Pereskioideae, or chains of fruits proliferating from fruit areoles (in Cylindropuntia fulgida), sessile (pedicellate in Pereskia), arising from stem areole at apex or axil of tubercle, radially symmetric [bilaterally symmetric]; flower tube 0.2-15[-30] cm; perianth epigynous (perigynous in some Pereskia), deciduous or persistent on fruit; tepals 5-50 or more, intergrading gradually from bractlike or sepal-like outer tepals to petal-like inner tepals; stamens usually 50-1500+ [sometimes fewer], decurrent on inner surface of flower tube; true ovary sunken in stem with tubercles present or absent, areoles conspicuous to obscure or absent; subtending scales persistent or deciduous, sometimes absent; spines present or absent, glochids present only in subfam. Opuntioideae; pistils compound, 1-locular; placentas parietal, 3-14[-20+]; style 1; nectary usually forming chamber around base of style; stigma lobes 3-14[-20+], 1 per placenta. Fruits basically berrylike (variable in succulence), deciduous or long persistent, indehiscent or dehiscent, succulent or leathery, sometimes promptly drying. Seeds (0-)5-3000+, yellowish, reddish, brown, black, or appearing tan or whitish (dark testa completely covered by pale, tough, glabrous or rarely pubescent, tight-fitting aril or "funicular envelope" in subfam. Opuntioideae), pyriform, obovoid, lenticular-reniform, or nearly circular, 0.4-12 mm diam.; testa glossy or dull; rarely with corky arillate appendages (strophioles in Mammillaria tetrancistra). x = 11.

Arbustes, arbres ou herbes pérennes, parfois épiphytes, rarement lactifères ; tiges simples ou ramifiées, souvent ± côtelées ou aplaties, succulentes, munies de mamelons souvent sur des hélices phyllotaxiques portant des aréoles munies de poils, d’aiguillons ou d’épines. Stipules absentes. Feuilles absentes ou réduites à des écailles, rarement développées, alternes, simples, succulentes ; limbe penninervé. Inflorescences terminant les aréoles ou à l’aisselle des mamelons, cymeuses, à fleurs en petits glomérules ou solitaires. Fleurs bisexuées, spirocycliques (aux pièces disposées en spirale), actinomorphes ou un peu zygomorphes, souvent sessiles et voyantes ; périanthe avec une transition graduelle des sépales en pétales ; sépales 3 à 6, peu développés ; pétales 5 à ∞, unis à la base et formant un hypanthe ; étamines peu à ∞, souvent ± soudées à la base des pétales, anthères introrses, s’ouvrant par des fentes longitudinales ; disque présent ; ovaire infère, avec 3 à ∞ carpelles, généralement uniloculaire, avec 15 à 100 ovules et placentation pariétale ; style 1, stigmates 3 à ∞. Fruits bacciformes, à nombreuses graines. Graines parfois arillées, à testa dur ou mou, exalbuminées.

Famille avec environ 90 à 140 genres (dépendant de l’auteur et de l’opinion de leur démarcation) et 1400 à 2000 espèces, tous les genres sauf un, en Amérique Centrale et du Sud ; le genre Rhipsalis s’étend en Afrique, à Madagascar et à Sri Lanka, avec une seule espèce au Gabon.

Arbustes'ou arbres, parfois épiphytes; tiges simples ou ramifiées, cylindriques et souvent ± côtelées ou aplaties, succulentes, vertes, munies de mamelons épars portant des aréoles poilues, aiguillonnées ou pourvues d'épines.'Feuilles'absentes ou réduites à des écailles, rarement développées, alternes et succulentes.'Fleurs'terminant les aréoles ou à l'aisselle des mamelons, solitaires ou en petits glomérules, souvent sessiles et voyantes, spirocycliques, ☿, actinomorphes ou un peu zygomorphes, généralement épigynes et à réceptacle pourvu d'un nectaire diversement conformé; périanthe avec transition graduelle des sépales en pétales; sépales 3-6, peu développés; pétales 5-∞, généralement libres; étamines peu-∞, souvent ± soudées à la base des pétales; gynécée à 3-∞ carpelles syncarpes; style avec stigmates en nombre variable; ovaire enfoncé dans le creux d'un réceptacle glabre, pourvu d'aréoles ou d'écailles, 1-loculaire, pariétal et à placentas ± saillants; ovules ∞, campylotropes, à funicules filiformes, allongés et libres ou soudés en faisceaux.'Fruits'bacciformes.'Graines ± nombreuses, parfois arillées, à testa dur ou mou, exalbuminées.\n\t\t\tFamille comprenant quelque 200 genres et plus de 2000 espèces répandues dans les régions arides de l'Amérique subtropicale et tropicale. Seul le genre'Rhipsalis's'étend jusque dans les tropiques de l'Ancien Monde: en Afrique, à Madagascar et à Ceylan. Certains'Opuntia , tel'O. ficus-indica (L.) Mill. ou figue de l'Inde, à baies comestibles, sont naturalisés dans toutes les régions subtropicales et tropicales.

Seeds mostly without endosperm Stamens numerous, free or adnate to the base of the petals Ovary inferior, 1-celled, with parietal many- or rarely few-ovuled placentas; stigma usually radiate Fruit baccate, many-seeded Succulent herbs and shrubs of diverse habit, often very spiny, and usually with much reduced leaves Flowers hermaphrodite, actinomorphic, often handsome, but very small in the African species (Rhipsalis) Calyx-tube adnate to the ovary and often produced beyond; lobes few to many, or reduced to minute teeth Petals 6 or more Stamens numerous, free or adnate to the base of the petals Ovary inferior, 1-celled, with parietal many- or rarely few-ovuled placentas; stigma usually radiate Fruit baccate, many-seeded Succulent herbs and shrubs of diverse habit, often very spiny, and usually with much reduced leaves Flowers hermaphrodite, actinomorphic, often handsome, but very small in the African species (Rhipsalis) Calyx-tube adnate to the ovary and often produced beyond; lobes few to many, or reduced to minute teeth Petals 6 or more

Carpels 3–?, syncarpous; ovary inferior, unilocular with 3–? parietal placentas; ovules ?; style single with 3–? stigmatic lobes Fruit a berry Flowers solitary on areoles, sessile (except in Pereskia), bisexual, usually actinomorphic; perianth segments ? (5–13 in species of FZ area), imbricate in bud, with gradual transition between sepals and petals, fused below to form a tube (hypanthium) Stamens ?, inserted at base of perianth; anthers 2-thecous, splitting longitudinally Seeds ? Succulent perennials with stems of varied shape and bristles arising from complex axillary structures (areoles) Fruit a berry Flowers solitary on areoles, sessile (except in Pereskia), bisexual, usually actinomorphic; perianth segments ? (5–13 in species of FZ area), imbricate in bud, with gradual transition between sepals and petals, fused below to form a tube (hypanthium) Stamens ?, inserted at base of perianth; anthers 2-thecous, splitting longitudinally Seeds ? Succulent perennials with stems of varied shape and bristles arising from complex axillary structures (areoles)

Fls epigynous, mostly regular and perfect; tepals generally numerous and spirally arranged, all showy and petaloid, or often the outer ones more sepaloid, but not sharply differentiated into two types, all united below to form a hypanthium; stamens numerous, arising spirally or in groups from the hypanthium; gynoecium of 3–many carpels united to form a compound, inferior, unilocular ovary with a single style and as many radiating stigmas as carpels; ovules numerous on parietal placentas, mostly campylotropous; fr a dry, pulpy or juicy berry, seeds with straight or more often curved, dicotyledonous embryo and abundant to scanty or no perisperm; spiny (seldom unarmed) stem- succulents, producing betalains but not anthocyanins, usually with scarcely developed lvs, the spines commonly restricted to regularly arranged areoles (the modified nodes) that usually also have clusters of small, detachable bristles (glochids); fls sessile, mostly large and solitary at the areoles. 30+/1000+.

Fleshy perennials, shrubs, trees or vines, terrestrial or epiphytic. Stems jointed, terete, globose, flattened, or fluted, mostly leafless and variously spiny. Leaves alternate, flat or subulate to terete, vestigial, or entirely absent; spines, glochids (easily detached, small, bristlelike spines), and flowers always arising from cushionlike, axillary areoles (modified short shoots). Flowers solitary, sessile, rarely clustered and stalked (in Pereskia), bisexual, rarely unisexual, actinomorphic or occasionally zygomorphic. Receptacle tube (hypanthium or perianth tube) absent or short to elongate, naked or invested with leaflike bracts, scales, areoles, and hairs, bristles, or spines; perianth segments usually numerous, in a sepaloid to petaloid series. Stamens numerous, variously inserted in throat and tube; anthers 2-loculed, dehiscing longitudinally. Ovary (pericarpel) inferior, rarely superior, 1-loculed, with 3 to many parietal (rarely basal) placentas; ovules usually numerous; style 1; stigmas 2 to numerous, papillate, rarely 2-fid. Fruit juicy or dry, naked, scaly, hairy, bristly, or spiny, indehiscent or dehiscent, when juicy then pulp derived from often deliquescent funicles (except in Pereskia). Seeds usually numerous, often arillate or strophiolate; embryo curved or rarely straight; endosperm present or absent; cotyledons reduced or vestigial, rarely leaflike.

Stems terete, globular, flattened or fluted, mostly leafless and variously spiny; spines always arising from complex axillary structures (areoles) Highly specialized fleshy perennials of diverse habit Perianth segments ?, closely imbricated in bud, in a sepaloid to petaloid series, ± free or fused below to form a short or elongate tube (hypanthium) Flowers solitary or rarely clustered, sessile (except in Pereskia), almost always bisexual, usually regular Fruit a dry or juicy berry, often spiny, bristly or scaly Ovary almost always inferior, with 3–? parietal placentas; ovules ?; style single; stigma-lobes 3–? Stamens ?; filaments variously inserted on or at base of perianth; anthers 2-thecous, splitting longitudinally Seeds ?, variously arillate or carunculate, with or without endosperm; embryo straight or curved Highly specialized fleshy perennials of diverse habit Perianth segments ?, closely imbricated in bud, in a sepaloid to petaloid series, ± free or fused below to form a short or elongate tube (hypanthium) Flowers solitary or rarely clustered, sessile (except in Pereskia), almost always bisexual, usually regular Fruit a dry or juicy berry, often spiny, bristly or scaly Ovary almost always inferior, with 3–? parietal placentas; ovules ?; style single; stigma-lobes 3–? Stamens ?; filaments variously inserted on or at base of perianth; anthers 2-thecous, splitting longitudinally Seeds ?, variously arillate or carunculate, with or without endosperm; embryo straight or curved

A large, almost exclusively American family, several of whose members are grown as ornamentals in East Africa. The epiphytic genus >i>Rhipsalis is sometimes considered to be indigenous in Africa, and a number of >i>Opuntia species have become widely naturalized