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The Mesembryanthemaceae form a major and unique component of southern
Africa's arid land flora. Succulent leaves, bright shiny-petalled
flowers and hygrochastic fruit (capsules open when it rains) characterize
the group. Mesembs are extremely diverse, particularly so in the
Succulent Karoo Region, although they have a strong presence in
the Fynbos. Some of the more primitive groups are almost entirely
confined to Fynbos, suggesting that the Fynbos Biome may have acted
as a mesic refuge for the semisucculent members of this group. Mesembs
are the subjects of a huge trade in 'curiosity plants' among succulent
collectors. They display features not seen elsewhere in the plant
kingdom. The combination of minutism, mimicry and extreme succulence
accounts for much of the variation in form and bizarre shapes that
add to their appeal.
Description
Mesembs
are annual to perennial succulents, ranging from minute to compact
and tufted to prostrate to decumbent; herbs, dwarf shrubs, subshrubs
or shrubs, they are rarely spiny. The leaves are opposite, rarely
alternate, simple, free at the base to almost completely united
into single, conical to ± spherical bodies, often sheathing
the stem. Their shape may be flat to 3-angled to cylindrical. All
leaves may be similar (plant homophyllous) or different types of
leaf pairs may be present (plant heterophyllous), members of a leaf
pair may be similar (plant isophyllous) or dissimilar (plant anisophyllous),
heterocellular (with distinct differences in cell sizes) or homocellular
(cells of similar size), ephemeral to persistent, with or without
bladder cells, sometimes tuberculate or punctate. The flowers are
bisexual, regular, in cymose inflorescences, sometimes representing
the entire plant, to solitary. The calyx is composed of 4 to 6 lobes,
sometimes forming a short tube above the ovary. The corolla is composed
of many more or less linear petals of staminodial origin, either
free or shortly fused at the base. Stamens are numerous, often surrounded
by few to many filamentous staminodes. The floral nectaries are
either koilomorphic: represented by usually 4 or 5 shell-shaped
depressions (in Mesembryanthemoideae), or lophomorphic: raised into
a crested ring, continuous or broken into ± 5 crested glands
(in some Ruschioideae), or flat, in the more primitive Ruschioideae.
The ovary is inferior to half-inferior (in Mesembryanthemoideae),
(3, 4)5(-25)-locular; placentas ranging from axile through basal
to parietal; ovules usually many; stigmas usually as many as locules
(seed chambers), subulate to filiform, sometimes fused at the base,
sometimes plumose. The fruit is a loculicidal capsule that splits
open when dry or mostly when wetted by means of hygroscopic expanding
keels, the fruit is rarely schizocarpic (splits into separate carpels)
with winged or nut-like mericarps, or it is fleshy and indehiscent.
The seeds are more or less kidney-shaped, glossy to textured, black
to brown to pale cream; in the seed, the embryo is curved around
the perisperm (food reserve tissue).
Distribution
Mesembs
comprise 127 genera and about 1 700 species, and account for 63%
of southern Africa's succulent flora, and almost 10% of South Africa's
Flora. The family Mesembryanthemaceae is comparable in size to the
New World Cactaceae, also an arid area specialist group. Mesembs
occur mostly in the southwestern parts of the African continent,
from ± 16° N in Angola down to the Western Cape Province,
extending well into the east over the central plateau of South Africa
and reaching into Zimbabwe and Botswana. They dominate vegetation
in the Succulent Karoo Region. This winter rainfall desert constitutes
the world's only arid biodiversity 'hotspot' of the 25 recognized
globally. The family is almost entirely endemic to southern Africa
with only about 20 species occurring naturally elsewhere. Several
species of Delosperma are found from northern Tanzania up
to the Yemen and also in the Arabian Peninsula and Madagascar. A
few species of Mesembryanthemum occur around the Mediterranean
Sea, and most species of the genus Disphyma are found in
Australia, New Zealand and the adjacent islands. The genus Sarcozona
and some species of Carpobrotus are restricted to Australia,
but species of Ruschia and Lampranthus may not be
indigenous to Australasia. It is also not clear whether species
of Malephora and Carpobrotus are recent introductions
to South America, and the populations of Carpobrotus found
along the west coast of North America were certainly introduced
by man.
Name and History
The original name given to mesembs was Mesembrianthemum this name
was chosen to describe the opening of flowers at noon. The spelling
was later changed to Mesembryanthemum. Historically, the group of
plants generally referred to as Aizoaceae have enjoyed fluctuating
status in various systematic treatments. Although mesembs are most
often classified in Aizoaceae, many taxonomists, particularly in
South African herbaria, prefer to treat them as a separate family,
Mesembryanthemaceae.
Ecology
The uniqueness and success of the group is attributable to combinations
of specialized morphological features that enable survival in arid
and in semi-arid conditions. The mostly hygrochastic fruit capsules
with rain-dispersed seed are diverse in form and serve taxonomically
in the distinction between genera. Succulence and innovative water
storage and saving mechanisms contribute to their resilience to
drought. In contrast to their close relatives in Aizoaceae, mesemb
flowers are characterized by whorls of bright-coloured petals originating
through a "dedoublement" of staminodial primordia. Flowers
are mostly adapted to insect pollinators.
Mesembs show a wide array of succulent morphologies, and succulence
is an adaptation to survive seasonal drought in arid environments,
although different strategies are evident. In succulent plants,
water relations are optimized and prioritized against other factors
such as carbon gain. Many utilize CAM photosynthesis, or are able
to switch to CAM. Besides low and sporadic rainfall, other ecologically
important factors in the life of mesembs include high irradiation,
high temperatures, soil salinity and sandblasting. Low temperatures
are thought to be a limiting factor in the distribution of most
mesembs in the colder interior of southern Africa.
As mesembs are extremely popular amongst specialist succulent plant
collectors, a number of them are threatened due to their removal
from the wild e.g. Conophytum burgeri, as well as through the destruction
of their habitats. Global climate change is seen as the greatest
threat to the Succulent Karoo Region, due to the shifting of climatic
belts, and the consequent destruction of their mega-niche.
Economic and cultural value
Their economic importance includes traditional uses in soap making,
preserves, poultices and preparations with psychoactive medicinal
properties, notably kougoed a preparation from Sceletium
species. Carpobrotus edulis, known as the sour fig, is eaten
as dried fruit or jam in the Western Cape. In addition, the astringent
juice of Carpobrotus leaves is used medicinally for sore
throats, bluebottle stings and fungal infections.
In the Garden
The
main use of mesembs is as garden plants. Their attractive flowers
produce colourful displays in spring and early summer. The bokbaaivygie
(Dorotheanthus bellidiformis)
is a favourite, but many species of Lampranthus and Drosanthemum
with their perennial shrubby habits are used in water-wise gardening.
Species of Carpobrotus and Aptenia are used as garden
plants worldwide. Among those previously described in the Plant
of the Week series are Lampranthus
aureus, L.affinis, L.
multiradiatus and L.haworthii;
Drosanthemum
speciosum and D.floribundum;
Frithia humilis and
F.pulchra; Lithops
olivacea, L.optica
and Conophytum truncatum.
- References
Leistner, O.A. (ed.). 2000. Seed plants of southern Africa: families
and genera. Strelitzia 10. National Botanical Institute,
Pretoria.
- Smith, G.F., Chesselet, P., Van Jaarsveld, E.J., Hartmann, H.,
Hammer, S., Van Wyk, B-E., Burgoyne, P., Klak, C. & Kurzweil,
H. 1998. Mesembs of the world. Briza Publications, Pretoria.
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Author
Pascale Chesselet
Kirstenbosch Research Centre
March 2004
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