Stangeria

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Stangeria
Stangeria eriopus001.jpg
S. eriopus in coastal lowland forest, South Africa
CITES Appendix I (CITES) [2]
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnosperms
Division: Cycadophyta
Class: Cycadopsida
Order: Cycadales
Family: Stangeriaceae
Subfamily: Stangerioideae
Genus: Stangeria
T.Moore
Species:
S. eriopus
Binomial name
Stangeria eriopus
Stangeria eriopus distribution.png
Range of Stangeria eriopus
Synonyms

Stangeria katzeriRegel
Stangeria paradoxaT. MooreStangeria sanderianaJ.Schust.
Stangeria schizodonW.Bull
Lomaria eriopus Kunze , 1839 Lomaria coriacea Kunze , 1836

Contents

Stangeria eriopus is a cycad endemic to southern Africa. It is the sole species in the genus Stangeria, most closely related to the Australian genus Bowenia , with which it forms the family Stangeriaceae.

Description

Stangeria eriopus is a very long-lived, perennial, evergreen cycad. The stalked, feathered, fern-like leaves 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), 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. [3] 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 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

Underground tuber stem Stangeria lignotuber.jpg
Underground tuber stem

Stems are completely subterranean and the root tuber is shaped like a carrot, 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. [4] The most common biflavones in the leaves are amentoflavone and bilobetin. [5]

Cones

S. eriopus reaches maturity at 5–7 years of age, and has stalked cones as reproductive organs. As is typical of cycads, the species is dioecious, 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.

Distribution and habitat

It is native to a narrow coastal strip, some 800 kilometres in extent, in the KwaZulu-Cape and Maputaland coastal forest mosaics of South Africa and southern Mozambique. 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 / 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 where it infests their cycads. The pest has now been introduced to Florida, Hawaii, Hong Kong and the Cayman Islands, 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 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. [6]

Discovery

When Gustav Kunze discovered the first plants, he incorrectly designated them as ferns 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. [7]

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 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. Common names includes Natal grass cycad, Hottentot's head and Stangeria.

Cytology

The species has 2n = 16 chromosomes. The cladogram shows twelve metacentric, two submetacentric and two acrocentric chromosomes. [8]

Systematics and Taxonomy

The closest relative to Stangeria eriopus was thought to be the genus Bowenia , both placed in the family Stangeriaceae . Another candidate is the extinct Tertiary genus Eostangeria . [5]

However, molecular phylogenetic studies show that Stangeria is more closely related to the genus Ceratozamia or to Zamia and Microcycas than to Bowenia, implying that the Stangeriaceae are not a monophyletic group. [8] [9] [10]

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. [11]

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. [12]

Related Research Articles

<span class="mw-page-title-main">Sporangium</span> Enclosure in which spores are formed

A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cycle. Sporangia can produce spores by mitosis, but in nearly all land plants and many fungi, sporangia are the site of meiosis and produce genetically distinct haploid spores.

<span class="mw-page-title-main">Cycad</span> Division of naked seeded dioecious plants

Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or female. Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow very slowly and live very long. Because of their superficial resemblance, they are sometimes mistaken for palms or ferns, but they are not closely related to either group.

<i>Bowenia</i> Genus of cycads in the family Stangeriaceae

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. B. spectabilis grows in warm, wet, tropical rainforests, on protected slopes and near streams, primarily in the lowlands of the Wet Tropics Bioregion. However, it has a local form with serrate pinna margins that grows in rainforest, Acacia-dominated transition forest, and also Casuarina-dominated sclerophyll forest on the Atherton Tableland, where it is subject to periodic bushfire. B. serrulata grows in sclerophyll forest and transition forest close to the Tropic of Capricorn.

<i>Cycas</i> Genus of cycads in the family Cycadaceae

Cycas is a genus of cycad, and the only genus in the family Cycadaceae. About 113 species are accepted, which are native to the Asia-Pacific, East Africa and Madagascar. Cycas circinalis, a species endemic to India, was the first cycad species to be described in western literature, and is the type species of the genus. The best-known Cycas species is Cycas revoluta.

<span class="mw-page-title-main">Zamiaceae</span> Family of cycads

The Zamiaceae are a family of cycads that are superficially palm or fern-like. They are divided into two subfamilies with eight genera and about 150 species in the tropical and subtropical regions of Africa, Australia and North and South America.

<span class="mw-page-title-main">Sporophyll</span>

A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls bear either megasporangia and thus are called megasporophylls, or microsporangia and are called microsporophylls. The overlap of the prefixes and roots makes these terms a particularly confusing subset of botanical nomenclature.

<span class="mw-page-title-main">Bennettitales</span> Extinct order of seed plants

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

<i>Retrophyllum</i> Genus of conifers

Retrophyllum is a genus of conifers in the family Podocarpaceae. It contains five generally recognized extant species with a disjunct distribution in the Southern Hemisphere, found in Papuasia and also in South America. Retrophyllum are evergreen trees typically occurring in tropical rainforests and cloud forests.

<i>Encephalartos</i> Genus of cycads in the family Zamiaceae

Encephalartos is a genus of cycad native to Africa. Several species of Encephalartos are commonly referred to as bread trees, bread palms or kaffir bread, since a bread-like starchy food can be prepared from the centre of the stem. The genus name is derived from the Greek words en (within), kephalē (head), and artos (bread), referring to the use of the pith to make food. They are, in evolutionary terms, some of the most primitive living gymnosperms.

<i>Dioon</i> Genus of cycads in the family Zamiaceae

Dioon is a genus of cycads in the family Zamiaceae. It is native to Mexico and Central America. Their habitats include tropical forests, pine-oak forest, and dry hillsides, canyons and coastal dunes.

A strobilus is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.

Pleuromeia is an extinct genus of lycophytes related modern quillworts (Isoetes). Pleuromeia dominated vegetation during the Early Triassic all over Eurasia and elsewhere, in the aftermath of Permian–Triassic extinction event. During this period it often occurred in monospecific assemblages. Its sedimentary context in monospecific assemblages on immature paleosols, is evidence that it was an opportunistic pioneer plant that grew on mineral soils with little competition. It spread to high latitudes with greenhouse climatic conditions. Conifers reoccurred in the Early Anisian, followed by the cycads and pteridosperms during the Late Anisian.

<i>Cycas micronesica</i> Species of cycad

Cycas micronesica is a species of cycad found on the island of Yap in Micronesia, the Marianas islands of Guam and Rota, and The Republic of Palau. It is commonly known as federico nut or fadang in Chamorro. The species, previously lumped with Cycas rumphii and Cycas circinalis, was described in 1994 by Ken Hill. Paleoecological studies have determined that C. micronesica has been present on the island of Guam for about 9,000 years. It is linked with Lytico-Bodig disease, a condition similar to amyotrophic lateral sclerosis (ALS), due to the neurotoxin BMAA found in its seeds, which were a traditional food source on Guam until the 1960s. The neurotoxin is present due to its symbiosis with cyanobacteria.

<i>Macrozamia glaucophylla</i>

Macrozamia glaucophylla is a species of cycad from the genus Macrozamia and the family Zamiaceae. Endemic to New South Wales, Australia, this species has features that resembles palms, although both species are taxonomically quite different. The current population trend of Macrozamia glaucophylla is stable with 2,500 to 10,000 mature individuals. The species are found in several habitats including forest and savanna. Ecologically, Macrozamia glaucophylla lives in terrestrial system, a land-based community of organisms where the biotic and abiotic components interact in the given area.

<i>Macrozamia heteromera</i>

Macrozamia heteromera is a species of cycad in the family Zamiaceae initially discovered by Charles Moore in 1858 and is endemic to New South Wales, Australia. It can be found in the north-western region of New South Wales within the Warrumbungle mountains and further south west towards the Coonabarabran district. It is a low trunked cycad usually at a height below 1 metre and can be found in dry sclerophyll woodlands. M. heteromera can be distinguished from the rest of the Macrozamia genus by its mid-green, narrow, usually divided pinnae and divided seedling pinnae. It is a plant that has toxic seeds and leaves, a characteristic common to cycads. However, after proper preparation and procedure, the seeds are fine for consumption.

<i>Macrozamia miquelii</i> Species of cycad

Macrozamia miquelii, is a species of cycad in the plant family Zamiaceae. It is endemic to Queensland and New South Wales in Eastern Australia. Located within sclerophyll forests dominated by eucalyptus trees, the cycad grows on nutrient-poor soils. It is recognised within the Zamiaceae family for its, medium height at 1 m, intermediate size of male and female cones and lighter green leaves compared to other cycads within the plant family of Zamiaceae. The seeds have an orange red sarcotesta which attracts fauna consumption, allowing a mutualistic seed dispersal for the cycad. These seeds are also edible for human consumption if prepared correctly to remove the toxins.

<i>Encephalartos ferox</i> Species of cycad

Encephalartos ferox, a member of the family Zamiaceae, is a small cycad with 35 cm wide subterranean trunk. It gets its name from the Latin word ferocious, likely from the spine-tipped lobes on the leaves of the plant. It is found naturally on the south-eastern coast of Africa where it has been used by local people for its starch content. It is considered to be one of the most popular cultivated cycads.

<i>Cycas pectinata</i> Species of cycad

Cycas pectinata was the fourth species of Cycas to be named; it was described in 1826 by Scottish surgeon and botanist Francis Buchanan-Hamilton from Kamrup, Assam in northeast India. The species is one of the most widespread cycads. It is found in the northeastern part of India, Nepal, Bhutan, northern Burma, southern China (Yunnan), Bangladesh, Burma, Malaysia, Cambodia, northern Thailand, Laos, and Vietnam. Cycas pectinata usually grow at elevation 300 m to 1200 m and in difficult terrains. In China, it grows in dry, open thickets in limestone mountains, red soil in sparse monsoon forests. Cycas pectinata grows up to 40 feet (12 m) tall and has very large, ovoid male cones. The tallest Cycas pectinata is a female plant in North Kamrup, Assam which measures 52.8 feet (16.1 m). The tree is the world's tallest Cycas plant. In Northeast India, the species is under severe threat due to clearing of forest and overcollection of male cones for preparation of traditional medicines. The species is listed in CITES Appendix II and IUCN Redlist.

<i>Cycas thouarsii</i> Species of evergreen plant

Cycas thouarsii, the Madagascar cycad, is an evergreen arborescent cycad in the genus Cycas. It is named after a French botanist Louis-Marie Aubert du Petit-Thouars (1758—1831).

<i>Zamia integrifolia</i> Species of cycad

Zamia integrifolia, also known as coontie palm is a small, tough, woody cycad native to the southeastern United States, the Bahamas, Cuba, and the Cayman Islands.

References

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  2. "Appendices | CITES". cites.org. Retrieved 2022-01-14.
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  7. Histoire des Plantes Monographie des Conifères, Gnétacées, Cycadacées, Alismacées, Triuridacées, Typhacées, Najadacées et Centrolépidacées. Paris, 1892, S. 68 (PDF)
  8. 1 2 Goro Kokubugata; Ken D. Hill; Katsuhiko Kondo (January 2002). abstract "Ribosomal DNA distribution in somatic chromosomes of Stangeria eriopus (Stangeriaceae, Cycadales) and molecular-cytotaxonomic relationships to some other cycad genera". Brittonia. 54 (1): 1–5. doi:10.1663/0007-196X(2002)054[0001:RDDISC]2.0.CO;2. S2CID   41822239.{{cite journal}}: Check |url= value (help)
  9. Rai, Hardeep S.; O’Brien, Heath E.; Reeves, Patrick A.; Olmstead, Richard G.; Graham, Sean W. (2003-11-01). "Inference of higher-order relationships in the cycads from a large chloroplast data set". Molecular Phylogenetics and Evolution. 29 (2): 350–359. doi:10.1016/S1055-7903(03)00131-3. ISSN   1055-7903. PMID   13678689.
  10. Salas-Leiva, Dayana E.; Meerow, Alan W.; Calonje, Michael; Griffith, M. Patrick; Francisco-Ortega, Javier; Nakamura, Kyoko; Stevenson, Dennis W.; Lewis, Carl E.; Namoff, Sandra (2013-11-01). "Phylogeny of the cycads based on multiple single-copy nuclear genes: congruence of concatenated parsimony, likelihood and species tree inference methods". Annals of Botany. 112 (7): 1263–1278. doi: 10.1093/aob/mct192 . ISSN   0305-7364. PMC   3806525 . PMID   23997230.
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  12. J. Keirungi; C. Fabricius (2005). "Selecting medicinal plants for cultivation at Nqabara on the Eastern Cape Wild Coast, South Africa". South African Journal of Science. 101 (11 & 12): 497–501. ISSN   0038-2353.