Crassulaceae J.St.-Hil.

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition

Characters of similar families: Saxifragaceae: leaves may be leathery but rarely succulent, leaves usually in a basal rosette, ovary inferior or semi-inferior, carpels 2 or rarely 3, fruit a capsule. Aizoaceae: leaf blade flat, terete, or triquetrous (3-angled), occasionally scale-like. Petals linear or absent, 0–250(–300). Stamens 1–500(–700), fruits usually capsules, sometimes indehiscent berries or nut-like.

Morphology General Habit

Succulent perennial herbs, rarely annuals or biennials, some shrubs

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves opposite or alternate, rarely verticillate, sometimes in a basal rosette, simple (compound in Kalanchoe), margins usually entire sometimes toothed or lobed

Morphology Reproductive morphology Inflorescences

Inflorescence terminal or axillary, usually cymous, sometimes a raceme or spike, occasionally flowers solitary

Morphology Reproductive morphology Flowers

Flowers bisexual (sometimes unisexual in Rhodiola), actinomorphic, perianth and androecium hypogynous or weakly perigynous, nectaries as scales at base of carpels

Morphology Reproductive morphology Flowers Calyx

Calyx (3–)4–5(–20) sepals, free or united into tube, entire

Morphology Reproductive morphology Flowers Corolla

Petals same number as sepals, free or united into tube

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens as many or twice as many as petals, free or fused with corolla tube; anthers basifixed and dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium Carpels

Carpels superior or partly inferior, free or united at base, as many as petals

Morphology Reproductive morphology Fruits

Fruit usually whorls of follicles, membranous or coriaceous; seeds 1–20+ per carpel, small, brownish.

Distribution

36 genera (29 in temperate regions) and ca. 1,400 species, although there are only 7 genera if Sedum is expanded to include all of the genera in subfamily Sempervivoideae. Widely distributed across the northern and southern hemispheres.

Note

Succulent perennials or shrubs, rarely annuals or biennials. Inflorescence terminal or axillary, with actinomorphic flowers. Sepals and petals of equal number. Carpels as many as petals. Small seeds held in follicles.

Description Author

Richard Wilford

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Succulent annual or perennial herbs, sometimes suffrutescent; leaves opposite, simple or sometimes compound, usually very thick; stipules absent

Morphology Reproductive morphology Flowers

Flowers regular, perfect, usually in cymes or panicles; sepals 4 or 5, united below; corolla 4–5-lobed, the lobes spreading. Stamens as many as or twice as many as the corolla-lobes, the anthers dehiscent by longitudinal slits

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, with as many carpels as sepals, distinct or partly united, with a scale at the base of each; ovules numerous, in 2 series along the ventral suture of the carpel

Morphology Reproductive morphology Fruits

Fruit consisting of 4 or 5 1-celled follicles containing numerous small or minute seeds; seeds with fleshy endosperm; embryo terete, with short, obtuse cotyledons.

Distribution

A widespread family of at least 33 genera and more than 1,000 species, in this hemisphere best represented in Mexico.

Crassulaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:1. 1954

Morphology General Habit

Herbs and undershrubs, usually succulent; leaves opposite or alternate, exstipulate

Morphology Reproductive morphology Flowers

Flowers actinomorphic, hermaphrodite, usually cymose

Morphology Reproductive morphology Flowers Calyx

Sepals free or united into a tube, often 4 or 5

Morphology Reproductive morphology Flowers Corolla

Petals the same number as the sepals, free or variously connate, hypogynous

Morphology Reproductive morphology Flowers Androecium

Stamens as many or twice as many as the petals, if few then alternate with the petals, slightly perigynous; filaments free; anthers 2-celled, introrse, dehiscing longitudinally

Morphology Reproductive morphology Flowers Nectaries

Hypogynous scales present within the stamens

Morphology Reproductive morphology Flowers Gynoecium

Carpels superior, the same number as the petals free or united at the base, 1-celled; ovules many or rarely few, inserted on the adaxial suture; styles short or elongated

Morphology Reproductive morphology Fruits

Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side

Morphology Reproductive morphology Seeds

Seeds mostly minute, usually with fleshy endosperm; embryo straight

Distribution

Widely distributed in temperate and tropical regions, especially in mountainous districts, rare in South America and Australasia

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Perennial herbs, rarely annual or biennial, rarely small shrubs, usually stem and/or leaf succulents

Morphology Leaves

Leaves opposite or alternate, rarely verticillate, exstipulate, usually simple, usually glabrous, often glaucous, rarely with glands in leaf surface, sometimes pubescent, hairs either unicellular or multicellular and glandular - capitate or eglandular, margins usually entire, sometimes serrate, crenate or dentate, rarely coarsely lobed

Morphology Reproductive morphology Flowers

Flowers usually in terminal cymose inflorescences, less often in spikes or racemes or solitary in leaf-axils, with or without bracts, regular, hermaphrodite, rarely unisexual and dioecious, mostly (3-)4-5 (-± 30)- merous; sepals 4-5 (-6), free or united into tube, persistent; petals same number as sepals, free or variously connate; stamens hypogynous or epipetalous, as many as petals or twice as many, in one whorl (Crassula-lineage) or more usually two whorls (Sedum-lineage), frequently obdiplostemenous, with outer whorl alternate and free from petals and inner whorl adnate to petals; filaments free or adnate to petals; anthers dorsifixed, bithecous, introrse, opening by a longitudinal slit; nectaries scale-like and usually present between the stamens and carpels

Morphology Reproductive morphology Flowers Gynoecium Carpels

Carpels superior, equal in number to petals, free or slightly connate at base, unilocular; ovules (few-) many, inserted on adaxial suture, submarginal or proximally axile; styles short or elongated, stigmatic surface on inner side of apex

Morphology Reproductive morphology Fruits

Fruit usually of separate follicles, rarely a capsule, follicles membranous or leathery, often surrounded by persistent membranous corolla, opening on adaxial side

Morphology Reproductive morphology Seeds

Seed minute, glabrous, testa variously striate and sometimes ornamented with ridges or papillae; endosperm usually present and sparse; embryo straight.

Distribution

Largest genus is Sedum L. with about 500 species in total. Echeveria DC. is the largest in the Western hemisphere with c. 150 species. Sedum follows Echeveria with c. 110 spp. in the Neotropics. Crassula L. has c. 11 spp. in the Neotropics. Villadia Rose contains c. 10 in Peru.

Diagnostic

Crassula spp. are usually prostrate or aquatic herbs, typically with isostemonous flowers. Echeveria spp. have obovatefleshy leaves, usually arranged in a rosette, and lateral racemose inflorescences Sedum spp. have paniculate cymes; both genera have free or basally connate petals. There are several species of Kalanchoe that are cultivated, some naturalized; they are best recognized by flowers with (4) fused petals forming a tube, and many species have lobed, crenate or serrate margined leaves. A number of species of Kalanchoe, formally in the genus Bryophyllum, have plantlets on the leaf margins.

Note

Main areas of speciation include Africa (dry areas), Madagascar, Macaronesia and Mexico. Up to six subfamilies are recognized but following DNA analysis only two main lineages are seen clearly - the Crassula-lineage and the Sedum-lineage. Mostly plants of dry, rocky habitats, usually terrestrial but rarely epiphytic and very rarely aquatic. Number of genera: There are four native genera (Sedum, Echeveria, Crassula and Villadia) and one cultivated (Kalanchoe).

Crassulaceae, G.E. Wickens. Flora of Tropical East Africa. 1987

Morphology General Habit

Annual, biennial or perennial herbs, rarely small shrubs, usually succulent

Morphology Leaves

Leaves opposite or alternate, rarely verticillate, exstipulate

Morphology Reproductive morphology Flowers

Flowers usually in terminal cymose inflorescences, less often in spikes or racemes or solitary in the leaf-axils, with or without bracts, regular, hermaphrodite, rarely unisexual and dioecious, mostly (3–)4–5(–±30)-merous

Morphology Reproductive morphology Flowers Calyx

Sepals free or united into a tube, persistent

Morphology Reproductive morphology Flowers Corolla

Petals the same number as the sepals, free or variously connate

Morphology Reproductive morphology Flowers Androecium

Stamens hypogynous or epipetalous, as many as the petals or twice as many, and in 2 whorls, frequently obdiplostemonous; filaments free or adnate to the petals; anthers 2-thecous, introrse, opening by a longitudinal slit; scale-like nectaries usually present between the stamens and carpels

Morphology Reproductive morphology Flowers Gynoecium

Carpels superior, equal in number to the petals, free or slightly connate at the base, 1-locular; ovules (few–)many, inserted on the adaxial suture; style short or elongated, stigmatose on the inner side of the apex

Morphology Reproductive morphology Fruits

Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side

Morphology Reproductive morphology Seeds

Seeds minute; endosperm usually present; embryo straight

Crassulaceae, R. Fernandes. Flora Zambesiaca 7:1. 1983

Morphology Leaves

Leaves opposite, verticillate or alternate, usually simple, sometimes compound, exstipulate, often ± thick and fleshy-succulent

Morphology Reproductive morphology Flowers

Flowers (3)4-5(6-32)-merous, actinomorphic, usually bisexual, in axillary or terminal cymes often grouped in corymb- or panicle-like inflorescences, less often in racemes or spikes or solitary in the leaf-axil

Morphology Reproductive morphology Flowers Calyx

Sepals free or ±: united at the base, sometimes in a ± long tube

Morphology Reproductive morphology Flowers Corolla

Petals as many as the sepals, free or ± connate, hypogynous

Morphology Reproductive morphology Flowers Androecium

Stamens as many or twice as many as the petals; filaments free or more or less connate with the corolla-tube; anthers 2-celled, introrse, basifixed, dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior; carpels equal in number to the petals, free or united (up to the middle); ovules many or few to one in each carpel, anatropous, inserted on the adaxial suture; styles short, sometimes nearly absent or ± elongate; stigmas capitate

Morphology Reproductive morphology Fruits

Fruit follicular, membranous or leathery, opening along the free part of the adaxial suture

Morphology Reproductive morphology Seeds

Seeds mostly minute with smooth, rugose or tuberculate tegument and usually fleshy endosperm; embryo straight

Morphology Reproductive morphology Flowers Nectaries

Hypogynous scales (scale-like nectaries) usually present, small, applied to the base of the outer face of the carpels