Scadoxus

{{Short description|Genus of flowering plants}} {{Automatic taxobox |image = Vegetation around Victoria falls.jpg |image_caption = ''Scadoxus multiflorus'' in Zambia |taxon = Scadoxus |authority = Raf.<ref name="WCSP_Scadoxus"/> |synonyms_ref = <ref name=WCSP_Scadoxus/> |synonyms = * ''Nerissa'' Salisb. * ''Demeusea'' De Wild. & T.Durand. * ''Choananthus'' Rendle |type_species = ''Scadoxus multiflorus'' |type_species_authority = Raf. }}

'''''Scadoxus''''' is a genus of African and Arabian plants in the Amaryllis family, subfamily Amaryllidoideae.<ref name=APweb/> 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.

thumb|''S. multiflorus'' (Blood Lily)

==Description==

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.<ref name=eMon_288235/>{{sfn|ps=|Hutchinson|2007}}

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.<ref name=eMon_288235/>{{sfn|ps=|Hutchinson|2014}}

==Taxonomy==

The genus was given its name in 1838 by Constantine Samuel Rafinesque. Rafinesque glossed the name as "umb. glor."<ref name=Rafi38/> (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.<ref name=pa/> The prefix ''sca'' may be derived from the Greek ''skia'' meaning "shade"<ref name=Pucc/> (''sciadon'' is the Greek equivalent of the Latin ''umbella'', "umbrella", used of flower heads in the form of umbels<ref name=Stea04/>).<ref name=Coom94/>

===Relationships===

''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'':<ref name=MeerClay04/>

{{clade |label1=Haemantheae |1={{clade |1={{clade |1=''Apodolirion'', ''Gethyllis'' |2={{clade |1=''Haemanthus'' |2='''''Scadoxus''''' }} }} |2={{clade |1=''Clivia'' |2=''Cryptostephanus'' }} }} }}

''Scadoxus'' was originally separated from ''Haemanthus'' by Rafinesque in 1838.<ref name=Rafi38/> His type species, ''Scadoxus multiflorus'', had been described as ''Haemanthus multiflorus'' by Thomas Martyn in 1795.<ref name=WCSP_Sm/> 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 2''n'' = 16 chromosomes, whereas ''Scadoxus'' species are found throughout tropical Africa, do not all form bulbs and have 2''n'' = 18 chromosomes.<ref name=MeerClay04/> 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.<ref name=NordDunc84/>

===Species=== {{As of|2014|March}}, the World Checklist of Selected Plant Families accepts the following nine species, one with three subspecies:<ref name=WCSP_Scadoxus/> {| class="wikitable" |- ! Image !! Scientific name !! Distribution |- |120px ||''Scadoxus cinnabarinus'' (Decne.) Friis & Nordal ||west and central Africa |- |120px ||''Scadoxus cyrtanthiflorus'' (C.H.Wright) Friis & Nordal ||Rwenzori Mountains in Uganda + Zaïre |- | ||''Scadoxus longifolius'' (De Wild. & T.Durand) Friis & Nordal || west Africa |- |120px ||''Scadoxus membranaceus'' (Baker) Friis & Nordal ||South Africa |- |120px ||''Scadoxus multiflorus'' (Martyn) Raf. (Blood Lily)||tropical and southern Africa and parts of the Arabian Peninsula |- | ||''Scadoxus nutans'' (Friis & I.Bjørnstad) Friis & Nordal ||Ethiopia |- |120px ||''Scadoxus pole-evansii'' (Oberm.) Friis & Nordal ||Zimbabwe |- | ||''Scadoxus pseudocaulus'' (I.Bjørnstad & Friis) Friis & Nordal ||tropical Africa |- |120px ||''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:<ref name=NordDunc84/> {{clade|style=line-height:100% |1={{clade |label1=rhizomes only |1={{cladex |1=''Scadoxus cyrtanthiflorus'' |2=''Scadoxus nutans'' |3={{clade |1=''Scadoxus pseudocaulus'' |2=''Scadoxus cinnabarinus'' }} }} |label2=bulbs |2={{cladex |1={{clade |1=''Scadoxus multiflorus'' |2=''Scadoxus pole-evansii'' }} |2={{clade |1=''Scadoxus puniceus'' |2=''Scadoxus membranaceus'' }} }} }} }} 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 and in the Arabian Peninsula. It has been introduced into parts of Mexico.<ref name=eMon_288235/> 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.<ref name=WCSP_309227/> By contrast, ''Scadoxus cyrtanthiflorus'' is found only in the Rwenzori Mountains on the border between Uganda and the Democratic Republic of the Congo.{{sfn|ps=|Hutchinson|2014}}

Most species are found in tropical forests, where they grow in warm, moist conditions in shade, either in soil or as epiphytes.{{sfn|ps=|Hutchinson|2014}} 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.{{sfn|ps=|Hutchinson|2014}}

==Cultivation==

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 {{convert|5|°C}}. ''S. multiflorus'' subsp. ''multiflorus'' is cultivated for sale in large quantities by the Dutch nursery industry.{{sfn|ps=|Hutchinson|2007}}

The tropical species require a minimum temperature of at least {{convert|10|°C}}, 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''.{{sfn|ps=|Hutchinson|2014}}

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.{{sfn|ps=|Hutchinson|2014}} In South Africa, where species such as ''S. puniceus'' can be grown outside, lily borer (''Brithys crini'') attacks plants.<ref name=pa/>

===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.{{sfn|ps=|Hutchinson|2007}}

==Toxicity and uses==

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.<ref name=pa_Sp/>

==References==

{{Reflist|refs= <ref name=APweb>Stevens, P.F. (2001 onwards), [http://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#AllAma "Asparagales: Amaryllidoideae"], ''Angiosperm Phylogeny Website'', retrieved 2014-02-25</ref>

<ref name=Coom94>{{Citation |last=Coombes |first=Allen J. |year=1994 |title=Dictionary of Plant Names |publication-place=London |publisher=Hamlyn Books |isbn=978-0-600-58187-1 }}, p. 167</ref>

<ref name=eMon_288235>{{Citation |title=''Scadoxus'' Raf. |work=eMonocot |url=http://e-monocot.org/taxon/urn:kew.org:wcs:taxon:288235 |access-date=2014-03-25 |archive-date=2014-01-23 |archive-url=https://web.archive.org/web/20140123064501/http://e-monocot.org/taxon/urn:kew.org:wcs:taxon:288235 |url-status=dead }}</ref>

<ref name=MeerClay04>{{Citation |last1=Meerow |first1=Alan W. |last2=Clayton |first2=Jason R. |year=2004 |title=Generic relationships among the baccate-fruited Amaryllidaceae (tribe Haemantheae) inferred from plastid and nuclear non-coding DNA sequences |journal=Plant Systematics and Evolution |volume=244 |issue=3–4 |pages=141–155 |doi=10.1007/s00606-003-0085-z |bibcode=2004PSyEv.244..141M |s2cid=10245220 |name-list-style=amp |url=https://naldc-legacy.nal.usda.gov/naldc/download.xhtml?id=2422&content=PDF |url-access=subscription }}{{dead link|date=June 2025|bot=medic}}{{cbignore|bot=medic}}</ref>

<ref name=NordDunc84>{{Citation |last1=Nordal |first1=I. |last2=Duncan |first2=T. |year=1984 |title=A cladistic analysis of ''Haemanthus'' and ''Scadoxus'' |journal=Nordic Journal of Botany |volume=4 |issue=2 |pages=145–153 |name-list-style=amp |doi=10.1111/j.1756-1051.1984.tb01482.x}}</ref>

<ref name=pa>{{Citation |title=''Scadoxus puniceus'' (L.) Friis & Nordal |work=PlantZAfrica |publisher=South African National Biodiversity Institute |url=http://www.plantzafrica.com/plantqrs/scadoxuspuniceus.htm |archive-url=https://web.archive.org/web/20110430041400/http://www.plantzafrica.com/plantqrs/scadoxuspuniceus.htm |archive-date=2011-04-30 |access-date=2014-03-25 }}</ref>

<ref name=pa_Sp>{{Citation |title=''Scadoxus multiflorus'' (Martyn) Raf. subsp. ''katharinae'' (Bak.) Friis & Nordal |work=PlantZAfrica |publisher=South African National Biodiversity Institute |url=http://www.plantzafrica.com/plantqrs/scadoxkath.htm |access-date=2014-03-25 }}</ref>

<ref name=Pucc>{{Citation |last=Puccio |first=Pietro |title=''Scadoxus multiflorus'' subsp. ''katharinae'' |work=photomazza.com |url=http://www.photomazza.com/?Scadoxus-multiflorus-subsp |access-date=2014-03-25 |others=transl. Mario Beltramini }}</ref>

<ref name=Rafi38>{{Citation |last=Rafinesque-Smaltz |first=C.S. |year=1838 |title=Flora Telluriana |publication-place=Philadelphia |url=https://www.biodiversitylibrary.org/page/7430162 |access-date=2014-03-25 |at=[https://www.biodiversitylibrary.org/page/7430513 part 4, p. 19]}}</ref>

<ref name=Stea04>{{Citation |last=Stearn |first=W.T. |year=2004 |title=Botanical Latin |publication-place=Newton Abbot |publisher=David & Charles |isbn=978-0-7153-1643-6 |page=271 }}</ref>

<ref name=WCSP_Scadoxus>{{Citation |contribution=Scadoxus|title=World Checklist of Selected Plant Families |publisher=Royal Botanic Gardens, Kew |contribution-url=http://apps.kew.org/wcsp/qsearch.do?plantName=Scadoxus |access-date=2014-03-23 }}</ref>

<ref name="WCSP_Sm">{{Citation |contribution=Scadoxus multiflorus|title=World Checklist of Selected Plant Families |publisher=Royal Botanic Gardens, Kew |contribution-url=http://apps.kew.org/wcsp/synonomy.do?name_id=287367 |access-date=2014-03-25 }}</ref>

<ref name=WCSP_309227>{{Citation |contribution=''Scadoxus multiflorus ''subsp.'' multiflorus''|title=World Checklist of Selected Plant Families |publisher=Royal Botanic Gardens, Kew |contribution-url=http://apps.kew.org/wcsp/namedetail.do?name_id=309227 |access-date=2014-03-25}}</ref> }}

==Bibliography== {{Wikispecies}} {{Commons category|Scadoxus}} *{{Citation |last=Duncan |first=G. |year=2013 |title=A synoptic review of ''Scadoxus'' Raf. (Amaryllidaceae) with notes on cultivation |journal=Herbertia |volume=66 |pages=189–224 }} *{{Citation |last=Hutchinson |first=J. |year=2007 |title=''Scadoxus'' of South Africa |journal=The Plantsman |series=New Series |volume=6 |issue=1 |pages=10–14 }} *{{Citation |last=Hutchinson |first=J. |year=2014 |title=''Scadoxus'' of central and east Africa |journal=The Plantsman |series=New Series |volume=13 |issue=1 |pages=36–42 }}

Category:Amaryllidaceae genera Category:Amaryllidoideae Category:Taxa named by Constantine Samuel Rafinesque