Lithops
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''Lithops'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
succulent plant In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', me ...
s in the ice plant family,
Aizoaceae The Aizoaceae, or fig-marigold family, is a large family of dicotyledonous flowering plants containing 135 genera and about 1800 species. They are commonly known as ice plants or carpet weeds. They are often called vygies in South Africa and N ...
. Members of the genus are native to
southern Africa Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number o ...
. The name is derived from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
words () 'stone' and () 'face', referring to the stone-like appearance of the plants. They avoid being eaten by blending in with surrounding rocks and are often known as pebble plants or living stones. The formation of the name from the Ancient Greek ''-ops'' means that even a single plant is called a Lithops.


Description

Individual ''Lithops'' plants consist of one or more pairs of bulbous, almost fused
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
opposite to each other and hardly any
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
. The slit between the leaves contains the
meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...
and produces
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s and new leaves. The leaves of ''Lithops'' are mostly buried below the surface of the
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
, with a partially or completely translucent top surface known as a leaf window which allows light to enter the interior of the leaves for
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
. During winter a new leaf pair, or occasionally more than one, grows inside the existing fused leaf pair. In
spring Spring(s) may refer to: Common uses * Spring (season), a season of the year * Spring (device), a mechanical device that stores energy * Spring (hydrology), a natural source of water * Spring (mathematics), a geometric surface in the shape of a h ...
the old leaf pair parts to reveal the new leaves and the old leaves will then dry up. ''Lithops'' leaves may shrink and disappear below ground level during
drought A drought is defined as drier than normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, an ...
. Lithops in habitat almost never have more than one leaf pair per head, presumably as an adaptation to the arid environment. Yellow or white flowers emerge from the fissure between the leaves after the new leaf pair has fully matured, one per leaf pair. This is usually in autumn, but can be before the summer
solstice A solstice is an event that occurs when the Sun appears to reach its most northerly or southerly excursion relative to the celestial equator on the celestial sphere. Two solstices occur annually, around June 21 and December 21. In many count ...
in '' L. pseudotruncatella'' and after the winter
solstice A solstice is an event that occurs when the Sun appears to reach its most northerly or southerly excursion relative to the celestial equator on the celestial sphere. Two solstices occur annually, around June 21 and December 21. In many count ...
in '' L. optica''. The flowers are often sweetly scented. The most startling
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
of ''Lithops'' is the colouring of the leaves. The leaves are
fenestrated A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical s ...
, and the epidermal windows are patterned in various shades of cream, grey, and brown, with darker windowed areas, dots, and red lines, according to species and local conditions. The markings function as remarkable
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
for the plant in its typical stone-like environment. As is typical of a window plant, the green tissue lines the inside of the leaves and is covered with
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable scattering of light. On a macroscopic scale (one in which the dimensions ...
tissue beneath the epidermal windows. ''Lithops'' are obligate outcrossers and require
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
from a separate plant. Like most mesembs, Lithops fruit is a dry capsule that opens when it becomes wet; some
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s may be ejected by falling raindrops, and the capsule re-closes when it dries out. Capsules may also sometimes detach and be distributed intact, or may disintegrate after several years.


Distribution

''Lithops'' occur naturally across wide areas of
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
, as well as small bordering areas in
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
and possibly
Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...
, from sea level to high mountains. Nearly a thousand individual populations are documented, each covering just a small area of dry grassland,
veld Veld ( or ), also spelled veldt, is a type of wide open rural landscape in :Southern Africa. Particularly, it is a flat area covered in grass or low scrub, especially in the countries of South Africa, Lesotho, Eswatini, Zimbabwe and Bot ...
, or bare rocky ground. Different ''Lithops'' species are preferentially found in particular environments, usually restricted to a particular type of rock. ''Lithops'' have not naturalised outside this region.
Rain Rain is water droplets that have condensed from atmospheric water vapor and then fall under gravity. Rain is a major component of the water cycle and is responsible for depositing most of the fresh water on the Earth. It provides water f ...
fall in ''Lithops'' habitats ranges from approximately 700 mm/year to near zero. Rainfall patterns range from exclusively summer rain to exclusively winter rain, with a few species relying almost entirely on dew formation for moisture. Temperatures are usually hot in summer and cool to cold in winter, but one species is found right at the coast with very moderate temperatures year round.


Cultivation

''Lithops'' are popular house plants and many specialist
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
growers maintain collections.
Seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s and plants are widely available in shops and over the
Internet The Internet (or internet) is the global system of interconnected computer networks that uses the Internet protocol suite (TCP/IP) to communicate between networks and devices. It is a '' network of networks'' that consists of private, p ...
. They are relatively easy to grow and care for if given sufficient sun and kept in well-draining soil. Normal treatment in mild
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
s is to keep them completely dry during winter, watering only when the old leaves have dried up and are replaced by a new leaf pair. Watering continues through autumn, when the plants flower, and then stops for winter. The best results are obtained in an environment with additional heat such as a greenhouse. In hotter
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
s, ''Lithops'' will have a summer dormancy when they should be kept mostly dry, and they may require some water in winter. In
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
climates, ''Lithops'' can be grown primarily in winter with a long summer dormancy. In all conditions, ''Lithops'' will be most active and need most water during autumn and each species will flower at approximately the same time. ''Lithops'' thrive best in a coarse, well-drained substrate. Any soil that retains too much water will cause the plants to burst their skins as they over-expand. Plants grown in strong light will develop hard strongly coloured skins which are resistant to damage and rot, although persistent overwatering will still be fatal. Excessive heat will kill potted plants as they cannot cool themselves by
transpiration Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. Water is necessary for plants but only a small amount of water taken up by the roots is used for growth ...
and rely on staying buried in cool soil below the surface. Commercial growers mix a mild fungicide or weak strength horticultural sulfur into the plant's water to prevent rotting. Lithops are sensitive to watering during hot weather, which can cause the plants to rot; in habitat the plants are often dormant when the temperatures are high, doing most of their growing during the cool months of the year. Low light levels will make the plants highly susceptible to rotting and fungal infection. In the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and ...
the following species have gained the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nor ...
’s
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
:- *'' Lithops karasmontana'' *'' Lithops olivacea'' *''
Lithops pseudotruncatella ''Lithops pseudotruncatella'' is a species of succulent in the family Aizoaceae. It is often confused with '' Conophytum truncatum,'' but with attention to detail, it can be distinguished. This confusion associated with ''L. pseudotruncatella'' i ...
'' *'' Lithops salicola'' *'' Lithops schwantesii''


Propagation

Propagation of ''Lithops'' is by
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
or cuttings. Cuttings can only be used to produce new plants after a plant has naturally divided to form multiple heads, so most propagation is by seed. ''Lithops'' can readily be pollinated by hand if two separate clones of a species flower at the same time, and seed will be ripe about 9 months later. Seed is easy to
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
, but the seedlings are small and vulnerable for the first year or two, and will not flower until at least two or three years old.


History

The first scientific description of ''Lithops'' was made by
botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
and artist William John Burchell, explorer of
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
, although he called it ''
Mesembryanthemum ''Mesembryanthemum'' is a genus of flowering plants in the family Aizoaceae; like many members of this family, it is characterized by long-lasting flower heads. Flowers of ''Mesembryanthemum'' protect their gametes from night-time dews or frosts ...
turbiniforme''. In 1811, Burchell discovered a specimen when picking up a "curiously shaped pebble" from the ground. Unfortunately the documented physical description was not detailed enough to be sure which ''Lithops'' he had discovered and the name ''Lithops turbiniformis'' is no longer used, although for many years it was applied to what is now known as '' Lithops hookeri''. Several more ''Lithops'' were published as ''
Mesembryanthemum ''Mesembryanthemum'' is a genus of flowering plants in the family Aizoaceae; like many members of this family, it is characterized by long-lasting flower heads. Flowers of ''Mesembryanthemum'' protect their gametes from night-time dews or frosts ...
''
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
until in 1922 N E Brown started to split up the overly large genus on the basis of the capsules. The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Lithops'' was created and dozens more species were published in the following decades. Brown, Gustav Schwantes, Kurt Dinter, Gert Nel, and Louisa Bolus continued to document ''Lithops'' from across southern Africa, but there was little consensus on the relationships between them, or even which populations should be grouped as species. As recently as the 1950s, the genus remained rather unknown in cultivation and was not well understood
taxonomically In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
. In the 1950s, Desmond and Naureen Cole began to study ''Lithops''. Together, the couple visited nearly all natural habitats of the different lithops populations and collected samples from approximately 400. They document and identify them, assigning a number, which is now known as the Cole number still used today all around the world. They studied and revised the genus, in 1988 publishing a definitive book (Lithops: Flowering Stones) describing the
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
,
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
, and varieties which have been accepted ever since. Because their camouflage is so effective, new species continue to be discovered, sometimes in remote regions of Namibia and South Africa, and sometimes in well-populated areas where they simply had been overlooked for generations. Recent discoveries include '' L. coleorum'' in 1994, '' L. hermetica'' in 2000, and '' L. amicorum'' in 2006.


Taxonomy

Many of the species listed have named
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
or varieties and some have many regional
form Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens. Form also refers to: *Form (document), a document (printed or electronic) with spaces in which to write or enter data * ...
s identified by old names or habitat locations. Identification of species is primarily by flower colour and leaf patterns.


Gallery

Lithops sp flower.jpg, ''Lithops'' sp. Blooms emerge between the leaves in autumn. Lithops sp Blooms.jpg, ''Lithops'' sp. Some species have flowers large enough to obscure the leaves. They open in the afternoon and close in the evening. Lithops sculpture.jpg, Sculpture of lithops, National Botanical Gardens of Ireland


References


Literature

* * * * *


External links


Lithops info from the BBC
(UK)
A guide to the cultivation of Lithops


an




Lithops photo album
(François Hoes, Belgium)
plantesdepedra.com
(Catalan info + translator)
manolithops.es
(Spanish info)
Scrapbooklithops
(General info & habitat photographs) {{Taxonbar, from=Q133283 Articles containing video clips Aizoaceae genera Taxa named by N. E. Brown Succulent plants