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Scadoxus
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| Scadoxus | |
|---|---|
| Scadoxus multiflorus in Zambia | |
| Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Monocots |
| Order: | Asparagales |
| Family: | Amaryllidaceae |
| Subfamily: | Amaryllidoideae |
| Genus: | Scadoxus Raf.[1] |
| Type species | |
| Scadoxus multiflorus | |
| Synonyms[1] | |
| |
Scadoxus is a genus of African and Arabian plants in the Amaryllis family, subfamily Amaryllidoideae.[2] The English names blood lily or blood flower are used for some of the species. The genus has close affinities with Haemanthus. Species of Scadoxus are grown as ornamental plants 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 alkaloids.

Description
[edit]Species of Scadoxus grow from bulbs or rhizomes. 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.[3][4]
The flowers are borne at the top of the scape in the form of a many-flowered umbel. Four or more bracts are present under the umbel at first. In some species, such as Scadoxus membranaceus, these bracts persist during flowering; in other species they wither before the flowers are fully open. Individual flowers have six red to pink tepals, joined at the base to form a tube. In most species, the flowers are more-or-less upright, although in Scadoxus cyrtanthiflorus the open flowers droop and in Scadoxus nutans the top of the scape bends over so that the flowers face downwards. The filaments of the stamens arise from the base of the tepals and may be flattened. The fruit takes the form of a globose berry, orange to red when ripe.[3][5]
Taxonomy
[edit]The genus was given its name in 1838 by Constantine Samuel Rafinesque. Rafinesque glossed the name as "umb. glor."[6] (possibly meaning umbella gloriosa, "glorious umbel"). Doxus, meaning "glory" or "splendour" in Greek, is usually interpreted as a reference to the often scarlet flowers of the genus.[7] The prefix sca may be derived from the Greek skia meaning "shade"[8] (sciadon is the Greek equivalent of the Latin umbella, "umbrella", used of flower heads in the form of umbels[9]).[10]
Relationships
[edit]Scadoxus is placed in tribe Haemantheae within the subfamily Amaryllidoideae, a tribe reserved for genera with fruit in the form of berries (baccate fruit). The tribe is predominantly African in origin and comprises six genera: Apodolirion, Gethyllis, Haemanthus, Scadoxus, Clivia and Cryptostephanus. The single most parsimonious phylogenetic tree found by analysis of both nuclear and plastid DNA in a 2004 study showed that Scadoxus is most closely related to Haemanthus:[11]
| Haemantheae |
| ||||||||||||||||||||||||
Scadoxus was originally separated from Haemanthus by Rafinesque in 1838.[6] His type species, Scadoxus multiflorus, had been described as Haemanthus multiflorus by Thomas Martyn in 1795.[12] 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 bulbs and have 2n = 16 chromosomes, whereas Scadoxus species are found throughout tropical Africa, do not all form bulbs and have 2n = 18 chromosomes.[11] 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.[13]
Species
[edit]As of March 2014[update], the World Checklist of Selected Plant Families accepts the following nine species, one with three subspecies:[1]
| Image | Scientific name | Distribution |
|---|---|---|
| Scadoxus cinnabarinus (Decne.) Friis & Nordal | west and central Africa | |
| Scadoxus cyrtanthiflorus (C.H.Wright) Friis & Nordal | Rwenzori Mountains in Uganda + Zaïre | |
| Scadoxus longifolius (De Wild. & T.Durand) Friis & Nordal | west Africa | |
| Scadoxus membranaceus (Baker) Friis & Nordal | South Africa | |
| Scadoxus multiflorus (Martyn) Raf. (Blood Lily) | tropical and southern Africa and parts of the Arabian Peninsula | |
| Scadoxus nutans (Friis & I.Bjørnstad) Friis & Nordal | Ethiopia | |
| Scadoxus pole-evansii (Oberm.) Friis & Nordal | Zimbabwe | |
| Scadoxus pseudocaulus (I.Bjørnstad & Friis) Friis & Nordal | tropical Africa | |
| Scadoxus puniceus (L.) Friis & Nordal | South Africa, Tanzania and Ethiopia |
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 was:[13]
| ||||||||||||||||||||||||||||||||||||||||
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
[edit]The genus as a whole is distributed in sub-Saharan Africa and in the Arabian Peninsula. It has been introduced into parts of Mexico.[3] The most widely distributed species is Scadoxus multiflorus, 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.[14] By contrast, Scadoxus cyrtanthiflorus is found only in the Rwenzori Mountains on the border between Uganda and the Democratic Republic of the Congo.[5]
Most species are found in tropical forests, where they grow in warm, moist conditions in shade, either in soil or as epiphytes.[5] The three species found in temperate regions of South Africa are more bulbous in habit than the tropical species; Scadoxus puniceus has been found growing in sand dunes and dry cliff faces.[5]
Cultivation
[edit]The cultivation of Scadoxus species has been described by Jonathan Hutchinson, the UK National Plant Collection 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 5 °C (41 °F). S. multiflorus subsp. multiflorus is cultivated for sale in large quantities by the Dutch nursery industry.[4]
The tropical species require a minimum temperature of at least 10 °C (50 °F), 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.[5]
Pests of Scadoxus in cultivation in the UK include mealy bugs and narcissus bulb fly (Merodon equestris). Red spider mites (Tetranychus species), slugs and snails can also cause problems.[5] In South Africa, where species such as S. puniceus can be grown outside, lily borer (Brithys crini) attacks plants.[7]
Cultivars
[edit]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.[4]
Toxicity and uses
[edit]The genus Scadoxus is known to have some strongly toxic species, containing poisonous alkaloids. 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 poisons and fishing poisons. Both species, as well as Scadoxus puniceus in South Africa, are used in traditional medicine.[15]
References
[edit]- ^ a b c "Scadoxus", World Checklist of Selected Plant Families, Royal Botanic Gardens, Kew, retrieved 2014-03-23
- ^ Stevens, P.F. (2001 onwards), "Asparagales: Amaryllidoideae", Angiosperm Phylogeny Website, retrieved 2014-02-25
- ^ a b c "Scadoxus Raf.", eMonocot, archived from the original on 2014-01-23, retrieved 2014-03-25
- ^ a b c Hutchinson 2007
- ^ a b c d e f Hutchinson 2014
- ^ a b Rafinesque-Smaltz, C.S. (1838), Flora Telluriana, Philadelphia, part 4, p. 19, retrieved 2014-03-25
- ^ a b "Scadoxus puniceus (L.) Friis & Nordal", PlantZAfrica, South African National Biodiversity Institute, archived from the original on 2011-04-30, retrieved 2014-03-25
- ^ Puccio, Pietro, "Scadoxus multiflorus subsp. katharinae", photomazza.com, transl. Mario Beltramini, retrieved 2014-03-25
- ^ Stearn, W.T. (2004), Botanical Latin, Newton Abbot: David & Charles, p. 271, ISBN 978-0-7153-1643-6
- ^ Coombes, Allen J. (1994), Dictionary of Plant Names, London: Hamlyn Books, ISBN 978-0-600-58187-1, p. 167
- ^ a b Meerow, Alan W. & Clayton, Jason R. (2004), "Generic relationships among the baccate-fruited Amaryllidaceae (tribe Haemantheae) inferred from plastid and nuclear non-coding DNA sequences", Plant Systematics and Evolution, 244 (3–4): 141–155, Bibcode:2004PSyEv.244..141M, doi:10.1007/s00606-003-0085-z, S2CID 10245220[dead link]
- ^ "Scadoxus multiflorus", World Checklist of Selected Plant Families, Royal Botanic Gardens, Kew, retrieved 2014-03-25
- ^ a b Nordal, I. & Duncan, T. (1984), "A cladistic analysis of Haemanthus and Scadoxus", Nordic Journal of Botany, 4 (2): 145–153, doi:10.1111/j.1756-1051.1984.tb01482.x
- ^ "Scadoxus multiflorus subsp. multiflorus", World Checklist of Selected Plant Families, Royal Botanic Gardens, Kew, retrieved 2014-03-25
- ^ "Scadoxus multiflorus (Martyn) Raf. subsp. katharinae (Bak.) Friis & Nordal", PlantZAfrica, South African National Biodiversity Institute, retrieved 2014-03-25
Bibliography
[edit]- Duncan, G. (2013), "A synoptic review of Scadoxus Raf. (Amaryllidaceae) with notes on cultivation", Herbertia, 66: 189–224
- Hutchinson, J. (2007), "Scadoxus of South Africa", The Plantsman, New Series, 6 (1): 10–14
- Hutchinson, J. (2014), "Scadoxus of central and east Africa", The Plantsman, New Series, 13 (1): 36–42
Scadoxus
View on GrokipediaTaxonomy
Etymology and history
The genus name Scadoxus was coined by the American naturalist Constantine Samuel Rafinesque in 1838, derived from Greek roots combining skiadion (meaning "umbel" or "parasol") and doxa (meaning "glory" or "splendor"), alluding to the striking, umbel-like inflorescences of its species.[7] Rafinesque glossed the name as "umb. glor.," interpreted as "glorious umbel," in his original description published in Flora Telluriana.[8] Rafinesque established the genus by separating it from Haemanthus, transferring Haemanthus multiflorus Martyn (now Scadoxus multiflorus) as the type species, based on differences in inflorescence structure and fruit characteristics.[8] This initial classification was later formalized within the Amaryllidaceae family by George Bentham and Joseph Dalton Hooker in their Genera Plantarum (volume 3, 1883), where Scadoxus was recognized alongside related African genera. Significant revisions occurred in the 1970s and 1980s through the work of botanists Ib Friis and Inger Nordal, who conducted detailed morphological studies on Haemanthus and reinstated Scadoxus as a distinct genus, transferring additional species such as Haemanthus puniceus L. (now Scadoxus puniceus) based on chromosome numbers, seed morphology, and vegetative traits.[9] Their key publication in 1976 divided Haemanthus sensu lato into Haemanthus s.s. and Scadoxus, enhancing taxonomic coherence.[10] As of updates in Plants of the World Online (POWO) reflecting taxonomic consensus through 2023, the genus comprises 9 accepted species, reflecting ongoing refinements from Friis and Nordal's framework without major alterations since the 1980s.[8]Phylogenetic relationships
Scadoxus is placed within the family Amaryllidaceae, subfamily Amaryllidoideae, and tribe Haemantheae, an entirely African clade characterized by baccate fruits and geophytic habits.[11] This positioning reflects the family's diversification in Africa during the Miocene, with Haemantheae representing one of its core tribes alongside others like Amaryllideae and Cyrtantheae.[12] Molecular phylogenetic studies using nuclear ribosomal ITS and plastid trnL-F sequences have confirmed the monophyly of Scadoxus within Haemantheae, with strong support from Bayesian inference (posterior probability 1.0) and moderate parsimony bootstrap values (86%).[13] The genus is most closely related to Haemanthus, forming a well-supported sister clade that shares bulbous storage organs and umbellate inflorescences with prominent stamens; this relationship is further corroborated by analyses of additional plastid markers such as rps16 and psbA-trnH.[14] In broader tribal analyses, Haemantheae allies with groups like Apodolirion and Gethyllis, underscoring the tribe's cohesion as a monophyletic African lineage.[15] Divergence time estimates for Haemantheae, based on calibrated molecular clocks, place the crown age at approximately 22 million years ago (95% highest posterior density: 15.2–29.9 Ma), aligning with Miocene climatic shifts that promoted diversification in southern and tropical African refugia.[12] Within Scadoxus, phylogenetic evidence highlights informal groupings based on underground habits, with some species exhibiting rhizomatous storage adapted to shaded, humid environments and others bulbous forms suited to seasonal dryness, though no formal subgenera are accepted due to the genus's polymorphic nature.[15]Accepted species
The genus Scadoxus comprises nine accepted species, all native to sub-Saharan Africa and the Arabian Peninsula.[8] These species exhibit variation in growth form, with some forming bulbs and others rhizomes, reflecting phylogenetic clustering into distinct groups.[8] The accepted species are as follows:- Scadoxus cinnabarinus (Decne.) Friis & Nordal: A rhizomatous species from West and Central Africa, noted for its scarlet flowers and evergreen foliage reaching up to 60 cm in height.[16][17]
- Scadoxus cyrtanthiflorus (C.H. Wright) Friis & Nordal: Bulbous, endemic to the Rwenzori Mountains in Central Africa, distinguished by its curved peduncle and 20–30 pendulous red flowers on stems up to 60 cm tall.[18][17]
- Scadoxus longifolius (De Wild. & T. Durand) Friis & Nordal: Bulbous geophyte from the Democratic Republic of Congo, featuring long, strap-shaped leaves and growing in wet tropical forests.[19][17]
- Scadoxus membranaceus (Baker) Friis & Nordal: The smallest species, bulbous and native to the Eastern Cape and KwaZulu-Natal in South Africa, with thin, membranous leaves and a height of about 40 cm; it is often evergreen in coastal habitats.[20][17]
- Scadoxus multiflorus (Martyn) Raf.: A widespread bulbous species across tropical Africa and Arabia, known as the blood lily; it includes subspecies such as S. m. subsp. multiflorus (deciduous, flowering without leaves in dry savannas) and S. m. subsp. katharinae (with deep pinkish-red flowers in coastal forests, often showing little dormancy).[21][17] Synonyms include Haemanthus multiflorus Martyn, transferred to Scadoxus by Rafinesque in 1838.[21]
- Scadoxus nutans (Friis & I.Bjørnstad) Friis & Nordal: Endemic to southwestern Ethiopia, a bulbous geophyte or epiphyte in montane forests, with nodding inflorescences of red flowers on stems up to 50 cm.[22][17]
- Scadoxus pole-evansii (Oberm.) Friis & Nordal: A rare bulbous species from southern Africa, particularly eastern Zimbabwe, with dark salmon-red flowers in groups of 50–70 on stems up to 130 cm; it is assessed as Near Threatened due to habitat loss in afromontane forests.[23][17]
- Scadoxus pseudocaulus (I. Bjørnstad & Friis) Friis & Nordal: Bulbous and evergreen, from southern Nigeria to west-central tropical Africa, forming a pseudostem up to 50 cm tall in shaded understory.[24][17]
- Scadoxus puniceus (L.) Friis & Nordal: Bulbous, known as the paintbrush lily, distributed in South and East Africa; it produces fluffy scarlet flower heads up to 15 cm across on stems to 75 cm, with a winter dormancy period.[25][17]
