Stapelieae
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Stapeliinae is a subtribe of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s within the tribe Ceropegieae of the subfamily
Asclepiadoideae The Asclepiadoideae are a subfamily of plants in the family Apocynaceae. Formerly, it was treated as a separate family under the name Asclepiadaceae, e.g. by APG II, and known as the milkweed family. They form a group of perennial herbs, twinin ...
of the family
Apocynaceae Apocynaceae (, from '' Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison. Notable mem ...
. The subtribe comprises about 35
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, including both the stem-succulent "stapeliads" and the horticulturally popular genera '' Brachystelma'' and '' Ceropegia''. The largest number of genera are native to Africa, but a more limited number of genera are widespread in Arabia and Asia. Historically, a similarly circumscribed taxon was treated as a separate tribe, Stapelieae.


Description

Most of the
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
of plants within the subtribe Stapeliinae are to varying degrees stem succulents, often referred to as "stapeliads". Many of these resemble
cacti A cactus (: cacti, cactuses, or less commonly, cactus) is a member of the plant family Cactaceae (), a family of the order Caryophyllales comprising about 127 genera with some 1,750 known species. The word ''cactus'' derives, through Latin, ...
, though are not closely related, as an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. The stems are often angular, mostly four-angled in cross-section, but in some species there are six or more, with some species of '' Hoodia'' having more than thirty angles. In size they vary from less than 2.5 cm/1" in length to over 2 m/6" tall. The leaves are in most species reduced to rudiments, sometimes hardened and thornlike, arranged on bumps or tubercles on the angles. Some species, however, still have recognisable leaves, most notably the Indian species '' Boucerosia frerei'' (syn. ''Frerea indica''), and some members of '' Tridentea''. Stapeliads are most abundant in warm, dry climates. In Africa, there are two separate regions where stapeliads have most diversified: northeast
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
, and
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
. Several species are endemic to the small island of
Socotra Socotra, locally known as Saqatri, is a Yemeni island in the Indian Ocean. Situated between the Guardafui Channel and the Arabian Sea, it lies near major shipping routes. Socotra is the largest of the six islands in the Socotra archipelago as ...
off the
Horn of Africa The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
. The
Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
, and most specifically the country of
Yemen Yemen, officially the Republic of Yemen, is a country in West Asia. Located in South Arabia, southern Arabia, it borders Saudi Arabia to Saudi Arabia–Yemen border, the north, Oman to Oman–Yemen border, the northeast, the south-eastern part ...
, contain another concentration of species. Several more are found in the drier parts of
Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...
,
Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan, is a landlocked country located at the crossroads of Central Asia and South Asia. It is bordered by Pakistan to the Durand Line, east and south, Iran to the Afghanistan–Iran borde ...
,
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Nepal Nepal, officially the Federal Democratic Republic of Nepal, is a landlocked country in South Asia. It is mainly situated in the Himalayas, but also includes parts of the Indo-Gangetic Plain. It borders the Tibet Autonomous Region of China Ch ...
, and
Myanmar Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and has ...
. A single species, '' Apteranthes europaea'' (syn. ''Caralluma europaea'') is found in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, in the very southern part of the
Iberian peninsula The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
and in the island of Lampedusa. Stapeliads are often regarded as a climax group within the family because of their often structurally complex
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s. Certain aspects of these reproductive parts mirror the
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
systems in the
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
family and represent a case of parallel
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
though both groups are quite unrelated and have developed similar, though not identical means to achieve the ultimate goal of pollination and therefore reproduction. Most stapeliads use flies as pollinators, that are attracted to odours resembling dung or rotting meat, emanating from the flowers. Many of the flowers also bear some physical resemblance to rotting animal carcasses, leading to their popular name of "carrion flowers". However, not all stapeliads smell bad, or attract flies. Some species use
beetles Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
,
bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
,
wasps A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
,
butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
or
moths Moths are a group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies (suborder Rhopalocera) a ...
as pollinators. Stapeliad flowers range in size from mere millimetres in species of '' Echidnopsis'' and '' Pseudolithos'' to those of ''
Stapelia gigantea ''Stapelia gigantea'' is a species of flowering plant in the genus '' Stapelia'' of the family Apocynaceae. Common names include Zulu giant, carrion plant and toad plant (although the nickname "carrion plant" can also refer to '' Stapelia grandif ...
'' that can reach 40 cm/16" in diameter, and are some the largest of flowers to be found on any species of succulent. Flowers are actinomorphic (radially symmetrical) with 5 free or fused sepals and petals. Some petals are formed in a shape of a star while some have a tubular shaped corolla. On some species, a raised ring or annulus is present in the center of the corolla. Instead of stamens and pistils, stapeliads contain a pollinarium.


Taxonomy

The first use of the subtribe (using the name Stapelieae) was by
George Don George Don (29 April 1798 – 25 February 1856) was a Scottish botanist and plant collector. Life and career George Don was born at Doo Hillock, Forfar, Angus, Scotland on 29 April 1798 to Caroline Clementina Stuart and George Don (b.1756), pr ...
in 1838. Don's taxon included both '' Ceropegia'' and ''
Stapelia ''Stapelia'' is a genus of low-growing, spineless, stem succulent plants, predominantly from South Africa with a few from other parts of Africa. Several Asian and Latin American species were formerly included but they have all now been transfer ...
''. Later workers, such as
George Bentham George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studie ...
, used these genera as the basis of two separate tribes, Ceropegieae and Stapeliae. The two tribes were formally reunited in 1991 as the tribe Ceropegieae. In 2004, Ulrich Meve and Sigrid Liede reintroduced the subtribe as a division of the tribe Ceropegieae, supported by
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies. Subsequent studies, including one in 2017, have confirmed the division of the tribe Ceropegieae into four subtribes, of which Stapeliinae is the most derived:


Genera

In 2014, Endress, Liede-Schumann and Meve placed the following genera in subtribe Stapeliinae. , some are treated as synonyms by
Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
. *''Anomalluma'' Plowes, synonym of '' Pseudolithos'' *'' Apteranthes'' J.C.Mikan *'' Australluma'' Plowes *'' Baynesia'' Bruyns *'' Boucerosia'' Wight & Arn. *'' Brachystelma'' Sims *''
Caralluma ''Caralluma'' is a genus of flowering plants in the family Apocynaceae, consisting of about 120 species. In 1795 William Roxburgh published the name '' Stapelia adscendens'' for a plant found in India. He commented that the name for the plant i ...
'' R.Br. *'' Caudanthera'' Plowes *'' Ceropegia'' L. *'' Desmidorchis'' Ehrenb. *'' Duvalia'' Haw. *'' Duvaliandra'' M.G.Gilbert *'' Echidnopsis'' Hook.f. *'' Edithcolea'' N.E.Br. *'' Hoodia'' Sweet ex Decne. *'' Huernia'' R.Br. *''
Larryleachia ''Larryleachia'' is a genus of stapeliad succulent flowering plants in the family Apocynaceae. ;Species * ''Larryleachia cactiformis'' (Hooker) Plowes * ''Larryleachia marlothii'' (N.E. Brown) Plowes * ''Larryleachia perlata'' (Dinter) Plowes ...
'' Plowes *'' Lavrania'' Plowes *'' Monolluma'' Plowes *'' Notechidnopsis'' Lavranos & Bleck *'' Ophionella'' Bruyns *'' Orbea'' Haw. *''Orbeanthus'' L.C.Leach, synonym of '' Orbea'' *'' Pectinaria'' Haw. *''
Piaranthus ''Piaranthus'' is a succulent plant genus in the subfamily Asclepiadoideae, in the family Apocynaceae. It was first described in 1810. Its name comes from Greek and is descriptive of the fleshy, succulent flowers typical of the genus (''"piar-" ...
'' R.Br. *'' Pseudolithos'' P.R.O.Bally *'' Quaqua'' N.E.Br. *'' Rhytidocaulon'' P.R.O.Bally *'' Richtersveldia'' Meve & Liede *'' Socotrella'' Bruyns & A.G.Miller *''
Stapelia ''Stapelia'' is a genus of low-growing, spineless, stem succulent plants, predominantly from South Africa with a few from other parts of Africa. Several Asian and Latin American species were formerly included but they have all now been transfer ...
'' L. *'' Stapelianthus'' Choux ex White & Sloane *'' Stapeliopsis'' Pillans *'' Tavaresia'' Welw. *'' Tridentea'' Haw. *''
Tromotriche ''Tromotriche'' is a genus of plant in family Apocynaceae. It is native to southern Africa.Gibbs Russell, G. E., W. G. M. Welman, E. Retief, K. L. Immelman, G. Germishuizen, B. J. Pienaar, M. Van Wyk & A. Nicholas. 1987. List of species of southe ...
'' Haw. *'' Whitesloanea'' Chiov. File:Baynesia lophophora.jpg, '' Baynesia lophophora'' File:Caralluma stalagmifera at Theni (17).jpg, '' Caralluma stalagmifera'' File:Ceropegia distincta.jpg, '' Ceropegia distincta'' File:Duvalia corderoyi.jpg, ''
Duvalia corderoyi ''Duvalia'' is a succulent plant genus in the subfamily Asclepiadoideae, in the family Apocynaceae (dogbane). The genus was first described in 1812, named after the French physician and botanist Henri-Auguste Duval (1777-1814). Description ...
'' File:Hoodia officinalis1MABJ.jpg, '' Hoodia officinalis'' File:Orbea_variegata(Stapelia_pulchellus).jpg, ''
Orbea variegata ''Orbea variegata'', also known as the star flower, carrion flower, starfish cactus and toad cactus, is a species of flowering plant in the family Apocynaceae that is native to the coastal belt of the Western Cape, South Africa, growing actively ...
'' File:Piaranthus gem glob.jpg, ''
Piaranthus geminatus ''Piaranthus'' is a succulent plant genus in the subfamily Asclepiadoideae, in the family Apocynaceae. It was first described in 1810. Its name comes from Greek and is descriptive of the fleshy, succulent flowers typical of the genus (''"piar-" ...
'' File:Quaqua mammillaris (Apocynaceae Asclepiadoideae) (28386411287).jpg, '' Quaqua mammillaris'' File:StapeliaFlavopurpureaFlower.jpg, '' Stapelia flavopurpurea''


References

* and {{taxonbar, from=Q2332067 Asclepiadoideae Plant subtribes