Scadoxus Puniceus - Kirstenbosch Botanical Garden - 1
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''Scadoxus'' is a genus of
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 ...
n and
Arabia The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Pl ...
n plants in the Amaryllis family, subfamily
Amaryllidoideae Amaryllidoideae (Amaryllidaceae ''s.s.'', amaryllids) is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. The most recent APG classification, APG III, takes a broad view of the Amaryllidaceae, which then ...
. The English names blood lily or blood flower are used for some of the species. The genus has close affinities with ''
Haemanthus ''Haemanthus'' is a Southern African genus of flowering plants in the family Amaryllidaceae, subfamily Amaryllidoideae. Members of the genus are known as blood lily and paintbrush lily. There are some 22 known species, native to South Africa, B ...
.'' Species of ''Scadoxus'' are grown as
ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that ...
s for their brilliantly coloured flowers, either in containers or in the ground in frost-free climates. Although some species have been used in traditional medicine, they contain poisonous
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s.


Description

Species of ''Scadoxus'' grow from
bulb In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs dur ...
s or
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ...
s. Bulbous species usually also have distinct rhizomes. Particularly in the non-bulbous species, the petioles (leaf stalks) overlap to produce a false stem or pseudostem, which may be purple-spotted. The leaf blades are lanceolate to ovate with a thickened midrib. The leafless flowering stem ( scape) is also sometimes purple-spotted, and either appears from among the leaves or pushes through the side of the pseudostem. The flowers are borne at the top of the scape in the form of a many-flowered
umbel In botany, an umbel is an inflorescence that consists of a number of short flower stalks (called pedicels) that spread from a common point, somewhat like umbrella ribs. The word was coined in botanical usage in the 1590s, from Latin ''umbella'' "p ...
. Four or more
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or o ...
s are present under the umbel at first. In some species, such as ''
Scadoxus membranaceus ''Scadoxus membranaceus'' is a flowering plant in the Amaryllidaceae family. It is a bulbous plant from South Africa (east Cape Province, KwaZulu-Natal). The smallest of the species of ''Scadoxus'', it is sometimes cultivated as an ornamental ...
'', these bracts persist during flowering; in other species they wither before the flowers are fully open. Individual flowers have six red to pink
tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s, joined at the base to form a tube. In most species, the flowers are more-or-less upright, although in ''
Scadoxus cyrtanthiflorus ''Scadoxus cyrtanthiflorus'' is a herbaceous plant endemic to the Rwenzori Mountains of east tropical Africa. Unusually for the genus ''Scadoxus'' its tubular blooms are pendant. It is sometimes grown as an ornamental plant in heated greenhouses ...
'' the open flowers droop and in ''
Scadoxus nutans ''Scadoxus nutans'' is a herbaceous plant endemic to southwest Ethiopia. Its red to pink flowers face downwards as the top of the flowering stem bends over, unlike any of the other species of ''Scadoxus''. It grows mainly as an epiphyte in tro ...
'' the top of the scape bends over so that the flowers face downwards. The filaments of the
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the fila ...
s arise from the base of the tepals and may be flattened. The fruit takes the form of a globose
berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, rasp ...
, orange to red when ripe.


Taxonomy

The genus was given its name in 1838 by
Constantine Samuel Rafinesque Constantine Samuel Rafinesque-Schmaltz (; October 22, 1783September 18, 1840) was a French 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ultimat ...
. Rafinesque glossed the name as "umb. glor." (possibly meaning ''umbella gloriosa'', "glorious umbel"). ''Doxus'', meaning "glory" or "splendour" in
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 ...
, is usually interpreted as a reference to the often scarlet flowers of the genus. The prefix ''sca'' may be derived from the Greek ''skia'' meaning "shade" (''sciadon'' is the Greek equivalent of the Latin ''umbella'', "umbrella", used of flower heads in the form of umbels).


Relationships

''Scadoxus'' is placed in tribe
Haemantheae Haemantheae are a tribe of subfamily Amaryllidoideae (family Amaryllidaceae). They are herbaceous monocot perennial flowering plants with a predominantly African distribution. Three subtribes are proposed and six genera including the type genus, ...
within the subfamily
Amaryllidoideae Amaryllidoideae (Amaryllidaceae ''s.s.'', amaryllids) is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. The most recent APG classification, APG III, takes a broad view of the Amaryllidaceae, which then ...
, a tribe reserved for genera with fruit in the form of
berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, ras ...
(baccate fruit). The tribe is predominantly African in origin and comprises six genera: ''
Apodolirion ''Apodolirion'' is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family (Amaryllidaceae, subfamily Amaryllidoideae). It consists of 6 species distributed in South Africa. The name ''Apodolirion'' comes from the Greek and m ...
'', ''
Gethyllis ''Gethyllis'' (probably from Greek ''"gethyon"'', bulb), commonly called Kukumakranka, Koekemakranka, or Kroekemakrank, is a genus of bulbous plant in the Amaryllid family with some 33 accepted species.. It is native to the Cape Provinces, t ...
'', ''
Haemanthus ''Haemanthus'' is a Southern African genus of flowering plants in the family Amaryllidaceae, subfamily Amaryllidoideae. Members of the genus are known as blood lily and paintbrush lily. There are some 22 known species, native to South Africa, B ...
'', ''Scadoxus'', '' Clivia'' and '' Cryptostephanus''. The single most parsimonious
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
found by analysis of both
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: *Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
and
plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyan ...
DNA in a 2004 study showed that ''Scadoxus'' is most closely related to ''Haemanthus'': ''Scadoxus'' was originally separated from ''Haemanthus'' by Rafinesque in 1838. His
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
, ''Scadoxus multiflorus'', had been described as ''Haemanthus multiflorus'' by Thomas Martyn in 1795. This separation was ignored by most workers until 1976, when ''Scadoxus'' was recognised as a distinct genus by Ib Friis and Inger Nordal. ''Haemanthus'' species are southern in distribution, form true
bulb In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs dur ...
s and have 2''n'' = 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 ar ...
s, whereas ''Scadoxus'' species are found throughout tropical Africa, do not all form bulbs and have 2''n'' = 18 chromosomes. The leaves of the two genera are also different. The leaves of ''Scadoxus'' species are thin, spirally arranged, with a distinct stalk ( petiole); in some species their bases form a pseudostem. The leaves of ''Haemanthus'' species are thicker, opposite, without a distinct petiole, and never form a pseudostem.


Species

, the
World Checklist of Selected Plant Families The World Checklist of Selected Plant Families (usually abbreviated to WCSP) is an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected pla ...
accepts the following nine species, one with three subspecies: Nordal and Duncan explored the relationship between eight of the species in a 1984 analysis based on morphological features (the little-known ''Scadoxus longiflorus'' was excluded). Their preferred
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
was: The main division is between a group which does not have bulbs, only rhizomes, and one which has bulbs, usually with rhizomes as well.


Distribution and habitat

The genus as a whole is distributed in
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
and in the
Arabian Peninsula The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plat ...
. It has been introduced into parts of
Mexico Mexico (Spanish language, Spanish: México), officially the United Mexican States, is a List of sovereign states, country in the southern portion of North America. It is borders of Mexico, bordered to the north by the United States; to the so ...
. The most widely distributed species is ''
Scadoxus multiflorus ''Scadoxus multiflorus'' (formerly ''Haemanthus multiflorus'') is a bulbous plant native to most of sub-Saharan Africa from Senegal to Somalia to South Africa. It is also native to Arabian Peninsula ( Saudi Arabia, Yemen, Oman) and to the S ...
,'' whose subspecies ''S. m.'' subsp. ''multiflorus'' is found throughout tropical and southern Africa and is the only member of the genus found in the Arabian Peninsula. By contrast, ''
Scadoxus cyrtanthiflorus ''Scadoxus cyrtanthiflorus'' is a herbaceous plant endemic to the Rwenzori Mountains of east tropical Africa. Unusually for the genus ''Scadoxus'' its tubular blooms are pendant. It is sometimes grown as an ornamental plant in heated greenhouses ...
'' is found only in the Rwenzori Mountains on the border between
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
and the
Democratic Republic of the Congo The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
. Most species are found in tropical forests, where they grow in warm, moist conditions in shade, either in soil or as
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s. The three species found in temperate regions of South Africa are more bulbous in habit than the tropical species; ''
Scadoxus puniceus ''Scadoxus puniceus'', commonly known as the paintbrush lily, is a species of bulbous plant. It is native to much of southern and eastern Africa: Ethiopia, Sudan, Tanzania, Malawi, Mozambique, Zambia, Zimbabwe, Botswana, Eswatini (Swaziland), ...
'' has been found growing in sand dunes and dry cliff faces.


Cultivation

The cultivation of ''Scadoxus'' species has been described by Jonathan Hutchinson, the UK
National Plant Collection The National Plant Collection scheme is the main conservation vehicle whereby the Plant Heritage charity (formerly the National Council for the Conservation of Plants and Gardens) can accomplish its mission: to conserve, grow, propagate, docume ...
holder for the genus. All species occur naturally in areas of summer rainfall, and in cultivation tend to start in growth in spring after a period of winter dormancy. The three species found in South Africa, ''S. multiflorus'', ''S. puniceus'' and ''S. membranaceous,'' are the most widely cultivated, being tolerant of winter temperatures down to . ''S. multiflorus'' subsp. ''multiflorus'' is cultivated for sale in large quantities by the Dutch nursery industry. The tropical species require a minimum temperature of at least , thriving in a warm, humid atmosphere. An open organic potting medium, similar to that used for orchids, suits all species. A coarse mix with considerable air spaces is particularly important for epiphytic species such as ''S. nutans''. Pests of ''Scadoxus'' in cultivation in the UK include
mealy bug In the theory of computation, a Mealy machine is a finite-state machine whose output values are determined both by its current state (computer science), state and the current inputs. This is in contrast to a Moore machine, whose output values are d ...
s and narcissus bulb fly (''
Merodon equestris ''Merodon equestris'' (Narcissus bulb fly, greater bulb fly, large bulb fly, large Narcissus fly) is a Holarctic species of hoverfly (Family Syrphidae). Like many other hoverflies it displays a colouration pattern similar to a stinging insect ...
''). Red spider mites ('' Tetranychus'' species), slugs and snails can also cause problems. In South Africa, where species such as ''S. puniceus'' can be grown outside, lily borer ('' Brithys crini'') attacks plants.


Cultivars

Some artificial hybrids between ''S. multiflorus'' subsp. ''katherinae'' and ''S. puniceus'' are known. Johannes Nicolai raised ''S.'' 'König Albert' which flowered for the first time in 1899. Although rare in cultivation, it multiplies rapidly. Of the same parentage is ''S.'' 'Andromeda', raised by C. G. van Tubergen around 1904.


Toxicity and uses

The genus ''Scadoxus'' is known to have some strongly toxic species, containing poisonous
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s. These are lethal to animals, such as sheep and goats, that graze on the leaves or bulbs. ''Scadoxus multiflorus'' and ''Scadoxus cinnabarinus'' are traditionally used in parts of tropical Africa as components of
arrow poison Arrow poisons are used to poison arrow heads or darts for the purposes of hunting and warfare. They have been used by indigenous peoples worldwide and are still in use in areas of South America, Africa and Asia. Notable examples are the poisons se ...
s and fishing poisons. Both species, as well as ''Scadoxus puniceus'' in South Africa, are used in traditional medicine.


References


Bibliography

* * * {{Taxonbar, from=Q142747 Amaryllidaceae genera Amaryllidoideae Taxa named by Constantine Samuel Rafinesque