Stangeria Eriopus
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''Stangeria eriopus'' is a
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
endemic to southern
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. It is the sole species in the genus ''Stangeria'', most closely related to the
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
n genus ''
Bowenia The genus ''Bowenia'' includes two living and two fossil species of cycads in the family Stangeriaceae, sometimes placed in their own family Boweniaceae. They are entirely restricted to Australia. The two living species occur in Queensland. '' ...
'', with which it forms the family
Stangeriaceae Stangeriaceae is the smallest family of the cycads, including only two extant genera, '' Stangeria'' and ''Bowenia'', which both have an underground stem. Taxonomy Although the family was first published by Schimper and Schenk as Stangerieae, t ...
.


Description

''Stangeria eriopus'' is a very long-lived, perennial, evergreen cycad. The stalked, feathered, fern-like
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are between 25 centimeters and two meters long, with the petiole comprising one third to one half of the overall length (in both varieties). They are pinnately-veined, which distinguishes the species from all other cycads. The petiole comprises half the length of the leaf. The young leaves are bent in bud position, the tip appears rolled up. Young leaves are dotted with short, gray hairs (
trichomes Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant ...
), which usually fall off quickly and only stick to the petiole. These trichomes are unbranched and transparent or colored. The species occurs as two variable forms or varieties. The forest form, growing in regions with higher rainfall, is characterized by large, wide leaves that can reach up to 2 m in length. The
grassland A grassland is an area where the vegetation is dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur natur ...
form, growing in regions subject to annual fire and drought, has shorter leaves with a thicker cuticle that may only be 30 cm long.


Tubers

Stems are completely subterranean and the root tuber is shaped like a
carrot The carrot ('' Daucus carota'' subsp. ''sativus'') is a root vegetable, typically orange in color, though purple, black, red, white, and yellow cultivars exist, all of which are domesticated forms of the wild carrot, ''Daucus carota'', nat ...
, which reaches a diameter of 10 to 25 centimeters. The tip bifurcates into several shoot tips. These form at the beginning wooly scales, but fall off early. As in other cycads, ''S. eriopus'' forms coralloid roots. These are specialized, plagiotropic (sideways-growing) roots housing colonies of cyanobacteria ''Bacillus radicola'' and ''Azotobacter sp.'' that fix nitrogen, much like the roots of legumes. The tuber is rich in carbohydrates and contains an exceptionally high concentration of sodium sulfate, which explains the breaking-irritant effect. The most common biflavones in the leaves are amentoflavone and bilobetin.


Cones

''S. eriopus'' reaches maturity at 5–7 years of age, and has stalked
cones A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
as reproductive organs. As is typical of cycads, the species is
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
, meaning that male and female cones are borne on different plants. Both male and female cones are pedunculated and covered with silvery hair when young, which is deciduous at maturity.


Male cone

The male cone is cylindrical and tapers towards the tip. At maturity reaches a diameter between 30 – 40 mm and becomes between 10 – 25 cm long. The cone then turns yellowish brown at maturity. The scales or microsporophyll, are arranged cylindrically around the pin axis. Their shape is triangular to rhomboid. Each microsporophyll forms about 150 pollen sacs, which are attached in groups of 3-6 at the bottom. At maturity, the pin axis extends and the scales are lifted apart and release the pollen.


Female cone

The female cones are ellipsoidal to conical with a rounded tip. The cones are about 18 inches long and reach a diameter of about 8 centimeters. At maturity, the cone turn to dark green. Like the male microsporophylls, the female megasporophylls are cylindrically arranged around the axis. Their tips form but six vertical lines. The ovules are formed at the base of megasporophyll and reach a size of 35 × 25 millimeters. When the cone is ripe, the megasporophylls separate. The micropyle exudes a drop of liquid that pulls the pollen capillary to the embryo sack when it dries. After ripening the seeds, the female cones dissolve and the seeds fall to the ground. They consist of a hard, dark red part, which is surrounded by a fleshy, purple seed coat, which later turns brown. They are about 2 inches long and about 14 millimeters wide.


Pollen

The cones are insect-pollinated, giving off a faint odor to attract beetle pollinators. At maturity they fall apart to reveal the seeds, which are 2–3 cm in length. The pollen carry on the outside a glycocalyx, which consists of densely packed cylindrical units measuring 20 to 150 nanometers. They are perpendicular to the plasma membrane. Below is the sporopollenin. The germinal openings (apertures) are more proximal than distal. At the distal end of each pollen grain, there is an unusual pit (pseudosulcus) that resembles a "failed" distal aperture. File:Stangeria eriopus002.jpg, File:Stangeria eriopus 2014-06-19 01.jpg, File:Stangeria eriopus, vroulike en manlike keëls, Ou Fort.jpg, File:Stangeria seed 005.jpg,


Distribution and habitat

It is native to a narrow coastal strip, some 800 kilometres in extent, in the KwaZulu-Cape and
Maputaland coastal forest mosaic The Maputaland coastal forest mosaic is a subtropical moist broadleaf forest ecoregion on the Indian Ocean coast of Southern Africa. It covers an area of in southern Mozambique, Eswatini, and the KwaZulu-Natal Province of South Africa. Mozamb ...
s 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 countri ...
and southern
Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi ...
. The western limit of the distribution area is located near Banjul in the district of Sarah Baartman. It is found within 50 km, but not closer than 2 or 3 km from the sea. This species of cycad is adaptable and is found in many habitats, from grassland to closed forest, whether in full sunshine or shade. The species has a low salt tolerance however. It is sometimes found in meadows near the coastal dunes, where the plants are protected from salt water. Sandy, slightly acidic soil is preferred, but at the northern limit of its range, ''Stangeria eriopus'' also grows on clay or very stony soils.


Conservation

IUCN Red List Category & Criteria: Vulnerable, mainly due to habitat loss and over-exploiting for traditional medicine. It is listed under
CITES Appendix I CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
/ EU Annex A, which prohibits international trade in specimens of this species except when the purpose of the import is not commercial, for instance for scientific research. The species is threatened by the destruction of its habitat and the unsustainable harvesting for traditional medicinal purposes. Another possible threat to the species is the lobster louse (Diaspididae) ''Aulacaspis yasumatsui''. The insect originally hails from
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
where it infests their cycads. The pest has now been introduced to Florida, Hawaii, Hong Kong and the
Cayman Islands The Cayman Islands () is a self-governing British Overseas Territory—the largest by population in the western Caribbean Sea. The territory comprises the three islands of Grand Cayman, Cayman Brac and Little Cayman, which are located to the ...
, where it causes significant damage to cycads. If the species is introduced to South Africa, it could drastically reduce or even destroy the Stangeria population in a short amount of time. Stangeria eriopus is also a carrier of the fungus '' Guignardia mangiferae'', which causes great damage to
citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
fruit, but remains on the plant without symptoms. The larvae of the butterfly '' Callioratis millari'' feed on the leaves of the species. The ''Stangeria eriopus'' can be asexually reproduced from root parts. It is the first species of cycad that has been propagated using tissue culture, which simplifies the conservation of the species.


Discovery

When Gustav Kunze discovered the first plants, he incorrectly designated them as
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s probably due to the primitive nerves, under the name ''Lomaria coriacea''. In 1839 he described it again as a separate species of fern as ''Lomaria eriopus''. It was not until 1851 that William Stanger discovered that they were cycads when he observed the cones. He sent samples to England, where they were described by Thomas Moore described the Art 1853 as ''Stangeria paradoxa'' and thus also established the genus. However, since the epithet "eriopus" of Kunze was validly described, Henri Ernest Baillon with his description in 1892 the correct name as ''Stangeria eriopus''.


Etymology

The genus was named in honor of William Stanger (1811-1854), who sent the first cones to England. He was an English physician and naturalist who worked in South Africa. The binomial name comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
prefix ''erio-'', meaning "woolly", and suffix ''-pus'', "footed", referring to the woolly petiole bases. It was named in honour of William Stanger, a former surveyor-general of
Natal NATAL or Natal may refer to: Places * Natal, Rio Grande do Norte, a city in Brazil * Natal, South Africa (disambiguation), a region in South Africa ** Natalia Republic, a former country (1839–1843) ** Colony of Natal, a former British colony ( ...
. Common names includes Natal grass cycad, Hottentot's head and Stangeria.


Cytology

The species has 2n = 16
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s. The cladogram shows twelve metacentric, two submetacentric and two acrocentric chromosomes.


Systematics and Taxonomy

The closest relative to ''Stangeria eriopus'' was thought to be the genus ''
Bowenia The genus ''Bowenia'' includes two living and two fossil species of cycads in the family Stangeriaceae, sometimes placed in their own family Boweniaceae. They are entirely restricted to Australia. The two living species occur in Queensland. '' ...
'', both placed in the family ''
Stangeriaceae Stangeriaceae is the smallest family of the cycads, including only two extant genera, '' Stangeria'' and ''Bowenia'', which both have an underground stem. Taxonomy Although the family was first published by Schimper and Schenk as Stangerieae, t ...
''. Another candidate is the extinct Tertiary genus '' Eostangeria''. However, molecular phylogenetic studies show that ''Stangeria'' is more closely related to the genus ''
Ceratozamia ''Ceratozamia'' is a genus of New World cycads in the family Zamiaceae. The genus contains 27 known currently living species and one or two fossil species. Most species are endemic to mountainous areas of Mexico, while few species extend into the ...
'' or to ''
Zamia ''Zamia'' is a genus of cycad of the family Zamiaceae, native to North America from the United States (in Georgia and Florida) throughout the West Indies, Central America, and South America as far south as Bolivia. The genus is considered to be ...
'' and ''
Microcycas ''Microcycas'' is a genus of cycads in the family Zamiaceae containing only one species, ''Microcycas calocoma'', endemic to a small area in western Cuba in Pinar del Río Province. Description The plant grows up to 10 m tall with an upright, so ...
'' than to ''Bowenia'', implying that the Stangeriaceae are not a monophyletic group.


Uses

In South African traditional medicine, the thickened subterranean tuber stem serves both for the production of various magical tinctures and as an emetic. The dried tuber is also mixed with feed to combat internal parasites in
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
. For these purposes, the plants are collected and sold, this goes so far that the stock is now endangered. For one gram of tuber, 5 cents were paid on the market in Mthala in 2005.


References

* * * * * *


External links

* *
The Cycad Pages: ''Stangeria eriopus''


{{Taxonbar, from=Q150991 Cycads Flora of Southern Africa Flora of South Africa Monotypic gymnosperm genera Near threatened flora of Africa