Many species of cactus have long, sharp spines, like this Opuntia.
A cactus (plural: cacti, cactuses, or cactus) is a member of the plant family Cactaceae,[Note 1] a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word "cactus" derives, through Latin, from the Ancient Greekκάκτος, kaktos, a name originally used by Theophrastus for a spiny plant whose identity is not certain. Cacti occur in a wide range of shapes and sizes. Most cacti live in habitats subject to at least some drought. Many live in extremely dry environments, even being found in the Atacama Desert, one of the driest places on earth. Cacti show many adaptations to conserve water. Almost all cacti are succulents, meaning they have thickened, fleshy parts adapted to store water. Unlike many other succulents, the stem is the only part of most cacti where this vital process takes place. Most species of cacti have lost true leaves, retaining only spines, which are highly modified leaves. As well as defending against herbivores, spines help prevent water loss by reducing air flow close to the cactus and providing some shade. In the absence of leaves, enlarged stems carry out photosynthesis. Cacti are native to the Americas, ranging from Patagonia in the south to parts of western Canada in the north—except for Rhipsalis baccifera, which also grows in Africa and Sri Lanka.
Cactus spines are produced from specialized structures called areoles, a kind of highly reduced branch. Areoles are an identifying feature of cacti. As well as spines, areoles give rise to flowers, which are usually tubular and multipetaled. Many cacti have short growing seasons and long dormancies, and are able to react quickly to any rainfall, helped by an extensive but relatively shallow root system that quickly absorbs any water reaching the ground surface. Cactus stems are often ribbed or fluted, which allows them to expand and contract easily for quick water absorption after rain, followed by long drought periods. Like other succulent plants, most cacti employ a special mechanism called "crassulacean acid metabolism" (CAM) as part of photosynthesis. Transpiration, during which carbon dioxide enters the plant and water escapes, does not take place during the day at the same time as photosynthesis, but instead occurs at night. The plant stores the carbon dioxide it takes in as malic acid, retaining it until daylight returns, and only then using it in photosynthesis. Because transpiration takes place during the cooler, more humid night hours, water loss is significantly reduced.
Many smaller cacti have globe-shaped stems, combining the highest possible volume for water storage, with the lowest possible surface area for water loss from transpiration. The tallest[Note 2] free-standing cactus is Pachycereus pringlei, with a maximum recorded height of 19.2 m (63 ft), and the smallest is Blossfeldia liliputiana, only about 1 cm (0.4 in) in diameter at maturity. A fully grown saguaro (Carnegiea gigantea) is said to be able to absorb as much as 200 U.S. gallons (760 l; 170 imp gal) of water during a rainstorm. A few species differ significantly in appearance from most of the family. At least superficially, plants of the genus Pereskia resemble other trees and shrubs growing around them. They have persistent leaves, and when older, bark-covered stems. Their areoles identify them as cacti, and in spite of their appearance, they, too, have many adaptations for water conservation. Pereskia is considered close to the ancestral species from which all cacti evolved. In tropical regions, other cacti grow as forest climbers and epiphytes (plants that grow on trees). Their stems are typically flattened, almost leaf-like in appearance, with fewer or even no spines, such as the well-known Christmas cactus or Thanksgiving cactus (in the genus Schlumbergera).
Cacti have a variety of uses: many species are used as ornamental plants, others are grown for fodder or forage, and others for food (particularly their fruit). Cochineal is the product of an insect that lives on some cacti.
The 1,500 to 1,800 species of cacti mostly fall into one of two groups of "core cacti": opuntias (subfamily Opuntioideae) and "cactoids" (subfamily Cactoideae). Most members of these two groups are easily recognizable as cacti. They have fleshy succulentstems that are major organs of photosynthesis. They have absent, small, or transient leaves. They have flowers with ovaries that lie below the sepals and petals, often deeply sunken into a fleshy receptacle (the part of the stem from which the flower parts grow). All cacti have areoles—highly specialized short shoots with extremely short internodes that produce spines, normal shoots, and flowers.
The remaining cacti fall into only two genera, Pereskia and Maihuenia, and are rather different, which means any description of cacti as a whole must frequently make exceptions for them. Pereskia species superficially resemble other tropical forest trees. When mature, they have woody stems that may be covered with bark and long-lasting leaves that provide the main means of photosynthesis. Their flowers may have superior ovaries (i.e., above the points of attachment of the sepals and petals), and areoles that produce further leaves. The two species of Maihuenia have small, globe-shaped bodies with prominent leaves at the top.
Cacti show a wide variety of growth habits, which are difficult to divide into clear, simple categories.
They can be tree-like (arborescent), meaning they typically have a single more-or-less woody trunk topped by several to many branches. In the genus Pereskia, the branches are covered with leaves, so the species of this genus may not be recognized as cacti. In most other cacti, the branches are more typically cactus-like, bare of leaves and bark, and covered with spines, as in Pachycereus pringlei or the larger opuntias. Some cacti may become tree-sized but without branches, such as larger specimens of Echinocactus platyacanthus. Cacti may also be described as shrubby, with several stems coming from the ground or from branches very low down, such as in Stenocereus thurberi.
Smaller cacti may be described as columnar. They consist of erect, cylinder-shaped stems, which may or may not branch, without a very clear division into trunk and branches. The boundary between columnar forms and tree-like or shrubby forms is difficult to define. Smaller and younger specimens of Cephalocereus senilis, for example, are columnar, whereas older and larger specimens may become tree-like. In some cases, the "columns" may be horizontal rather than vertical. Thus, Stenocereus eruca has stems growing along the ground, rooting at intervals.
Cacti whose stems are even smaller may be described as globular (or globose). They consist of shorter, more ball-shaped stems than columnar cacti. Globular cacti may be solitary, such as Ferocactus latispinus, or their stems may form clusters that can create large mounds. All or some stems in a cluster may share a common root.
Other cacti have a quite different appearance. In tropical regions, some grow as forest climbers and epiphytes. Their stems are typically flattened, almost leaf-like in appearance, with fewer or even no spines. Climbing cacti can be very large; a specimen of Hylocereus was reported as 100 meters (330 ft) long from root to the most distant stem. Epiphytic cacti, such as species of Rhipsalis or Schlumbergera, often hang downwards, forming dense clumps where they grow in trees high above the ground.
The leafless, spiny stem is the characteristic feature of the majority of cacti (and all of those belonging to the largest subfamily, the Cactoideae). The stem is typically succulent, meaning it is adapted to store water. The surface of the stem may be smooth (as in some species of Opuntia) or covered with protuberances of various kinds, which are usually called tubercles. These vary from small "bumps" to prominent, nipple-like shapes in the genus Mammillaria and outgrowths almost like leaves in Ariocarpus species. The stem may also be ribbed or fluted in shape. The prominence of these ribs depends on how much water the stem is storing: when full (up to 90% of the mass of a cactus may be water), the ribs may be almost invisible on the swollen stem, whereas when the cactus is short of water and the stems shrink, the ribs may be very visible.
The stems of most cacti are some shade of green, often bluish or brownish green. Such stems contain chlorophyll and are able to carry out photosynthesis; they also have stomata (small structures that can open and close to allow passage of gases). Cactus stems are often visibly waxy.
Flowers appear from the upper part of an areole, spines from the lower (Cereus species)
Areoles are structures unique to cacti. Although variable, they typically appear as woolly or hairy areas on the stems from which spines emerge. Flowers are also produced from areoles. In the genus Pereskia, believed similar to the ancestor of all cacti, the areoles occur in the axils of leaves (i.e. in the angle between the leaf stalk and the stem). In leafless cacti, areoles are often borne on raised areas on the stem where leaf bases would have been.
Areoles are highly specialized and very condensed shoots or branches. In a normal shoot, nodes bearing leaves or flowers would be separated by lengths of stem (internodes). In an areole, the nodes are so close together, they form a single structure. The areole may be circular, elongated into an oval shape, or even separated into two parts; the two parts may be visibly connected in some way (e.g. by a groove in the stem) or appear entirely separate (a dimorphic areole). The part nearer the top of the stem then produces flowers, the other part spines. Areoles often have multicellular hairs (trichomes) that give the areole a hairy or woolly appearance, sometimes of a distinct color such as yellow or brown.
In most cacti, the areoles produce new spines or flowers only for a few years, and then become inactive. This results in a relatively fixed number of spines, with flowers being produced only from the ends of stems, which are still growing and forming new areoles. In Pereskia, a genus close to the ancestor of cacti, areoles remain active for much longer; this is also the case in Opuntia and Neoraimondia.
The great majority of cacti have no visible leaves; photosynthesis takes place in the stems (which may be flattened and leaflike in some species). Exceptions occur in three groups of cacti. All the species of Pereskia are superficially like normal trees or shrubs and have numerous leaves. Many cacti in the opuntia group (subfamily Opuntioideae, opuntioids) also have visible leaves, which may be long-lasting (as in Pereskiopsis species) or be produced only during the growing season and then be lost (as in many species of Opuntia). The small genus Maihuenia also relies on leaves for photosynthesis. The structure of the leaves varies somewhat between these groups. Pereskia species have "normal" leaves, with a midrib and a flattened blade (lamina) on either side. Opuntioids and Maihuenia have leaves that appear to consist only of a midrib.
Even those cacti without visible photosynthetic leaves do usually have very small leaves, less than 0.5 mm (0.02 in) long in about half of the species studied and almost always less than 1.5 mm (0.06 in) long. The function of such leaves cannot be photosynthesis; a role in the production of plant hormones, such as auxin, and in defining axillary buds has been suggested.
Botanically, "spines" are distinguished from "thorns": spines are modified leaves, and thorns are modified branches. Cacti produce spines, always from areoles as noted above. Spines are present even in those cacti with leaves, such as Pereskia, Pereskiopsis and Maihuenia, so they clearly evolved before complete leaflessness. Some cacti only have spines when young, possibly only when seedlings. This is particularly true of tree-living cacti, such as Rhipsalis or Schlumbergera, but some ground-living cacti, such as Ariocarpus, also lack spines when mature.
The spines of cacti are often useful in identification, since they vary greatly between species in number, color, size, shape and hardness, as well as in whether all the spines produced by an areole are similar or whether they are of distinct kinds. Most spines are straight or at most slightly curved, and are described as hair-like, bristle-like, needle-like or awl-like, depending on their length and thickness. Some cacti have flattened spines (e.g. Schlerocactus papyracanthus). Other cacti have hooked spines. Sometimes, one or more central spines are hooked, while outer spines are straight (e.g., Mammillaria rekoi).
In addition to normal-length spines, members of the subfamily Opuntioideae have relatively short spines, called glochids, that are barbed along their length and easily shed. These enter the skin and are difficult to remove, causing long-lasting irritation.
Most ground-living cacti have only fine roots, which spread out around the base of the plant for varying distances, close to the surface. Some cacti have taproots; in genera such as Copiapoa, these are considerably larger and of a greater volume than the body. Taproots may aid in stabilizing the larger columnar cacti. Climbing, creeping and epiphytic cacti may have only adventitious roots, produced along the stems where these come into contact with a rooting medium.
Like their spines, cactus flowers are variable. Typically, the ovary is surrounded by material derived from stem or receptacle tissue, forming a structure called a pericarpel. Tissue derived from the petals and sepals continues the pericarpel, forming a composite tube—the whole may be called a floral tube, although strictly speaking only the part furthest from the base is floral in origin. The outside of the tubular structure often has areoles that produce wool and spines. Typically, the tube also has small scale-like bracts, which gradually change into sepal-like and then petal-like structures, so the sepals and petals cannot be clearly differentiated (and hence are often called "tepals"). Some cacti produce floral tubes without wool or spines (e.g. Gymnocalycium) or completely devoid of any external structures (e.g. Mammillaria). Unlike the flowers of other cacti, Pereskia flowers may be borne in clusters.
Cactus flowers usually have many stamens, but only a single style, which may branch at the end into more than one stigma. The stamens usually arise from all over the inner surface of the upper part of the floral tube, although in some cacti, the stamens are produced in one or more distinct "series" in more specific areas of the inside of the floral tube.
The flower as a whole is usually radially symmetrical (actinomorphic), but may be bilaterally symmetrical (zygomorphic) in some species. Flower colors range from white through yellow and red to magenta.