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{{short description|
{{Use Australian English|date=July 2024}}
{{Use dmy dates|date=September 2019}}
▲{{short description|A large shrub or small tree in the family Proteaceae found on the south coast of Western Australia}}
{{featured article}}
{{Speciesbox
|name = Showy banksia
|image = Banksia speciosa - San Francisco Botanical Garden.jpg
|image_caption =
|parent = Banksia ser. Banksia
|display_parents = 4
|
▲|binomial = ''Banksia speciosa''
▲|binomial_authority = [[Robert Brown (botanist, born 1773)|R.Br.]]<ref name=apni>{{APNI | name = ''Banksia speciosa'' R.Br. | id = 55431}}</ref>
|synonyms = ''Sirmuellera speciosa'' <small>(R.Br.) [[Otto Kuntze|Kuntze]]</small>
'''''Banksia speciosa''''', commonly known as the '''showy banksia''', is a species of large shrub or small tree in the [[family (biology)|family]] [[Proteaceae]]. It
Collected and described by [[Robert Brown (botanist, born 1773)|Robert Brown]] in the early 19th century, ''B. speciosa'' is classified in the [[Series (botany)|series]] ''[[Banksia ser. Banksia|Banksia]]'' within the genus. Its closest relative is ''[[Banksia baxteri|B. baxteri]]''. ''B. speciosa'' plants are killed by [[Bushfires in Australia|bushfire]], and regenerate from seed. The flowers attract nectar- and insect-feeding birds, particularly [[honeyeater]]s, and a variety of insects. In cultivation, ''B. speciosa'' grows well in a sunny location on well-drained soil in areas with dry summers. It cannot be grown in areas with humid summers, though it has been [[Grafting|grafted]] onto ''[[Banksia serrata]]'' or ''[[Banksia integrifolia|B. integrifolia]]''.
==Description==
[[File:Speciosagdn3.JPG|thumb
''B. speciosa'' grows as a shrub or small tree anywhere from 1 to 6 or rarely 8 m (4–26 ft) high.<ref name="FloraBase"/> It has an open many-branched [[Habit (biology)|habit]], arising from a single stem or trunk with smooth grey bark. Unlike many banksias, it does not have a [[lignotuber]].<ref name="George 1981">{{cite journal | last = George | first=Alex S. |
The cream to yellow flower spikes, known as [[inflorescence]]s, can appear at any time of year.<ref name="George 1981"/> They arise on the ends of one- or two-year-old stems and are roughly cylindrical in shape with a domed apex,<ref name="George 1981"/><ref name = "Wrigley 1991"/> measuring {{convert|4|–|12|cm|in|abbr=on}} high and {{convert|9|–|10|cm|in|abbr=on}} wide at [[anthesis]]. Each is a compound flowering structure, with a large number of individual flowers arising out of a central woody axis.<ref name="George 1981"/> A field study on the southern [[sandplain]]s revealed an average count of 1369±79 on each spike.<ref name="witkowski91">{{cite journal|last1 = Witkowski
The seed is {{convert|3.7|–|4.5|cm|in|abbr=on}} long and fairly flattened, and is composed of the seed body proper, measuring {{convert|1|–|1.4|cm|in|abbr=on|
A variant from the [[Gibson, Western Australia|Gibson]] area has an upright habit and leaves.<ref name=atlas/> Otherwise, ''B. speciosa'' shows little variation across its range. Combined with its vigour and prominence in its habitat, this has led George to speculate that it is a recent development among its relatives.<ref name="George 1981"/>
''[[Banksia baxteri]]'' resembles ''B. speciosa'' and co-occurs with it at the western edge of
==Taxonomy==
[[File:Banksia speciosa (Bauer).jpg|right|thumb|Watercolour by [[Ferdinand Bauer]] c. 1811, painted from the sketches he made on board the ''[[HMS Investigator (1798)|Investigator]]'', in the company of [[Robert Brown (botanist, born 1773)|Robert Brown]]]]
The first botanical collector of this species may well have been [[Claude Riche]], [[naturalist]] to [[Bruni d'Entrecasteaux]]'s 1791 expedition in search of the lost ships of [[Jean-François de Galaup, comte de La Pérouse]]. During a visit to [[Esperance Bay]], Riche explored an area in which ''B. speciosa'' is extremely common. However, he got lost and was forced to abandon his collections.<ref>{{cite book | last = Duyker | first = Edward |
Robert Brown recorded 31 species of ''Banksia'' in his 1810 work ''[[Prodromus Florae Novae Hollandiae et Insulae Van Diemen]]'', and in his [[Brown's taxonomic arrangement of Banksia|taxonomic arrangement]], placed the taxon in the [[subgenus]] ''[[Banksia verae]]'', the "True Banksias", because the inflorescence is a typical ''Banksia'' flower spike.<ref name="Brown 1810">{{cite book | last = Brown | first = Robert | year = 1810 | title = Prodromus Florae Novae Hollandiae et Insulae Van Diemen | publisher = Richard Taylor and Company | location = London, United Kingdom | language =
[[George Bentham]] published a thorough revision of ''Banksia'' in his landmark publication ''[[Flora Australiensis]]'' in 1870. In [[Bentham's taxonomic arrangement of Banksia|Bentham's arrangement]], the number of recognised ''Banksia'' species was reduced from 60 to 46. Bentham defined four [[Section (botany)|sections]] based on leaf, [[style (botany)|style]] and pollen-presenter characters. ''B. speciosa'' was placed in section ''Orthostylis''.<ref name="Bentham 1870">{{cite encyclopedia | last = Bentham | first = George |
In 1891, German botanist [[Otto Kuntze]] challenged the generic name ''Banksia'' [[Carolus Linnaeus the Younger|L.f.]], on the grounds that the name ''Banksia'' had previously been published in 1775 as ''Banksia'' [[Johann Reinhold Forster|J.R.Forst]] & [[Georg Forster|G.Forst]], referring to the genus now known as ''[[Pimelea]]''. Kuntze proposed ''Sirmuellera'' as an alternative, republishing ''B. speciosa'' as ''Sirmuellera speciosa''.<ref name="Kuntz1891">{{cite book|last=Kuntze|first=Otto|
===Current placement===
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Alex George published a new [[George's taxonomic arrangement of Banksia|taxonomic arrangement]] of ''Banksia'' in his classic 1981 monograph [[The genus Banksia L.f. (Proteaceae)|''The genus ''Banksia'' L.f. (Proteaceae)'']].<ref name="George 1981"/> Endlicher's ''Eubanksia'' became [[Banksia subg. Banksia|''B.'' subg. ''Banksia'']], and was divided into three sections. ''B. speciosa'' was placed in [[Banksia sect. Banksia|''B.'' sect. ''Banksia'']], and this was further divided into nine series, with ''B. speciosa'' placed in [[Banksia ser. Banksia|''B.'' ser. ''Banksia'']].
He thought its closest relative was clearly ''Banksia baxteri'' based on their similar appearance, noting the two overlapped in their
[[Kevin Thiele]] and [[Pauline Ladiges]] published a new arrangement for the genus in 1996; their [[morphology (biology)|morphological]] [[Cladistics|cladistic]] analysis yielded a [[cladogram]] significantly different from George's arrangement. [[Thiele and Ladiges' taxonomic arrangement of Banksia|Thiele and Ladiges' arrangement]] retained ''B. speciosa'' in series ''Banksia'', placing it in [[Banksia subser. Cratistylis|''B.'' subser. ''Cratistylis'']] along with ''B. baxteri'' as its [[Sister group|sister]] taxon and seven other Western Australian species.<ref name="Thiele 1996">{{cite journal | last1 = Thiele | first1 = Kevin |
:'''Genus ''[[Banksia]]'''''
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:::::''[[Banksia sceptrum|B. sceptrum]]''
In 2002, a [[molecular phylogenetics|molecular]] study by [[Austin Mast]] again showed ''B. speciosa'' and ''B. baxteri'' to be each other's closest relatives, but they were only distantly related to other members of the series ''Banksia''. Instead, their next closest relative turned out to be the distinctive ''[[Banksia coccinea]]''.<ref name="Mast 2002">{{cite journal | first1 = Austin R. | last1 = Mast |
Mast, Eric Jones and Shawn Havery published the results of their cladistic analyses of [[DNA sequence]] data for ''Banksia'' in 2005. They inferred a [[phylogeny]] greatly different from the accepted taxonomic arrangement, including finding ''Banksia'' to be [[paraphyly|paraphyletic]] with respect to ''[[Banksia ser. Dryandra|Dryandra]]''.<ref name="Mast 2005">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Jones | first2 = Eric H. | last3 = Havery | first3 = Shawn P. | year = 2005 | volume = 18 | issue = 1 | title = An Assessment of Old and New DNA Sequence Evidence for the Paraphyly of ''Banksia'' with Respect to ''Dryandra'' (Proteaceae) | journal = [[Australian Systematic Botany]] | pages = 75–88 | doi = 10.1071/SB04015}}</ref> A new taxonomic arrangement was not published at the time, but early in 2007 Mast and Thiele initiated a rearrangement by transferring ''Dryandra'' to ''Banksia'', and publishing [[Banksia subg. Spathulatae|''B.'' subg. ''Spathulatae'']] for the species having spoon-shaped cotyledons; in this way they also redefined the [[autonym (botany)|autonym]] ''B.'' subg. ''Banksia''. They foreshadowed publishing a full arrangement once DNA [[sampling (statistics)|sampling]] of ''Dryandra'' was complete. In the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then ''B. speciosa'' is placed in ''B.'' subg. ''Banksia''.<ref name="Mast 2007">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Thiele | first2 = Kevin | year = 2007 | title = The Transfer of ''Dryandra'' R.Br. to ''Banksia'' L.f. (Proteaceae) | journal = [[Australian Systematic Botany]] | volume = 20 | pages = 63–71 | doi = 10.1071/SB06016}}</ref>
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==Distribution and habitat==
[[File:Banksia speciosa map.png|thumb|right|Distribution of ''B. speciosa'' (showy banksia), shown on a map of Western Australia's biogeographic regions.<ref name="FloraBase"/>]]
''B. speciosa'' occurs on coastal dunes and sandplains in the [[Esperance Plains]] and [[Mallee (biogeographic region)|Mallee]] [[biogeography|biogeographic]] regions on the south coast of Western Australia,<ref name="FloraBase">{{FloraBase | name = ''Banksia speciosa'' R.Br
''B. speciosa'' grows on flat or gently sloping ground on deep white or grey sand. It is often the dominant shrub in shrubland, commonly found with such species as ''[[Lambertia inermis]]'',<ref name="George 1981"/> ''[[Banksia pulchella]]'', and ''[[Banksia petiolaris|B. petiolaris]]''.<ref name=atlas/>
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The prominent flower spikes are visited by many birds and insects. Honeyeaters are common visitors, particularly the [[New Holland honeyeater]], as well as the [[fuscous honeyeater]], [[western wattlebird]] and [[western spinebill]]. Other birds recorded foraging include the [[grey butcherbird]] and species of [[Acanthiza|thornbill]]. Insects recorded include ants, bees, wasps, butterflies, moths, flies and beetles.<ref name=atlas/> The [[short-billed black cockatoo]] breaks off old cones with follicles to eat the seed, often doing so before the seed is ripe.<ref name="witkowski91"/>
''B. speciosa'' is [[serotiny|serotinous]], that is, it has an [[Canopy seed bank|aerial seed bank]] in its [[Canopy (biology)|canopy]] in the form of the follicles of the old flower spikes. These are opened by fire and release seed in large numbers, which germinate and grow after rain. Seed can last for many years; old spikes 11 to 12 years old have been found to have 50% viable seed.<ref>{{cite journal|first1=Byron B. | last1=Lamont | first2=D. C. | last2=Le Maitre | first3=R. M. | last3=Cowling | first4=N. J. | last4=Enright|year=1991|title=Canopy Seed Storage in Woody Plants|journal=The Botanical Review|volume=57|issue=4|pages= 277–317 [284] |doi=10.1007/BF02858770| s2cid=37245625 }}</ref> Flower spikes appear to have similar numbers of follicles regardless of the age of the parent plant. Young plants begin flowering three years after regenerating from bushfire and store progressively larger numbers of old flowerheads (and hence seed) in the canopy. In one study, decade-old plants averaged around 3.5 old cones, whereas 21-year-old plants had 105, and were calculated as having over 900 viable seeds per plant. Plants appear to have a life span of at least 40 years, as healthy and vigorous individuals of this age are known.<ref name="witkowski91"/> An experimental burn and monitoring of resultant seedling germination and growth showed ''B. speciosa'' seeds, though numerous, had poor rates of establishment but that seedlings were able to access water more easily and had higher rates of survival after two years than co-occurring ''Banksia'' species. Though this suggested ''B. speciosa'' might outcompete its [[conspecific]]s, the authors of the study noted that there could be other factors not accounted for in its natural environment.<ref name="lamont 1995">{{cite journal|last1=Lamont | first1=Byron B. | last2=Witkowski | first2=E.T.F|year=1995|title=A Test for Lottery Recruitment among Four ''Banksia'' Species Based on Their Demography and Biological Attributes|journal=Oecologia|volume=101|issue=3|pages=299–308|jstor=4220888|doi=10.1007/BF00328815| pmid=28307050 | bibcode=1995Oecol.101..299L | s2cid=24306122 }}</ref>
''B. speciosa'' is extremely sensitive to dieback caused by ''[[Phytophthora cinnamomi]]'' and numbers in [[Cape Le Grand National Park|Cape Le Grand]] and [[Cape Arid National Park]]s have been drastically reduced as whole populations of plants have perished after exposure.<ref name=atlas/> It is an [[indicator species]] for the presence of the disease.<ref>{{cite web|url=http://www.dec.wa.gov.au/pdf/projects/dieback/dieback_indicators.pdf |title=Common Indicator Species for the Presence of Disease caused by ''Phytophthora cinnamomi'' |publisher=Government of Western Australia |
The tiny [[ascomycota|sac fungus]] ''[[Phyllachora banksiae]]'' subspecies ''westraliensis'' has been described from the leaves of ''B. speciosa'', its sole host. This fungus manifests as round flat cream-coloured spots around 1–3 mm in diameter on the upper leaf surface. The surrounding leaf tissue is sometimes discoloured orange. One or two shiny black fruit bodies measuring around 0.25–0.75 by 0.25–1 mm appear in the centre of the spots.<ref>{{cite journal|last1=Pearce | first1=Ceridwen A. | last2=Reddell | first2=Paul | last3=Hyde | first3=Kevin D.|year=2001|title=Revision of the Phyllochoraceae (Ascomycota) on Hosts in the Angiosperm Family Proteaceae|journal=Australian Systematic Botany|volume=14|issue=2|pages=283–328|doi=10.1071/SB00006}}</ref>
==Cultivation==
A fast-growing and attractive plant, ''B. speciosa'' grows readily in a sunny location in dry climates on well-drained soil, but does poorly in areas of humid summer climate, such as Australia's east coast. It has been [[Grafting|grafted]] successfully onto ''[[Banksia serrata]]'' and ''[[Banksia integrifolia|B. integrifolia]]'' to enable cultivation in these areas.<ref name = "Wrigley 1991"/><ref name=Collins>{{cite book | last1 = Collins | first1 = Kevin | last2 = Collins | first2 = Kathy | last3 = George | first3 = Alex S. | pages = 48, 98–99 |title = Banksias | publisher = Bloomings Books | location = Melbourne, Victoria | year = 2008 | isbn = 978-1-876473-68-6 }}</ref> Seeds do not require any treatment, and take 27 to 41 days to [[germinate]].<ref name="Sweedman 2006">{{cite book | last1=Sweedman | first1=Luke | last2=Merritt | first2=David | year = 2006 | title = Australian Seeds: A Guide to their Collection, Identification and Biology | url=https://archive.org/details/australianseedsg00auth | url-access=limited | publisher = CSIRO Publishing| location= Collingwood, Victoria | isbn = 0-643-09298-6 | page = [https://archive.org/details/australianseedsg00auth/page/n219 203]}}</ref> A specimen flowered in a greenhouse in the [[Royal Botanic Garden Edinburgh]] in 1830.<ref name="curtis"/> ''B. speciosa'' is an important [[cut flower]] crop.<ref name=Collins/> It was one of several species considered for commercial cropping in [[Tenerife]], and trials showed that seedlings were moderately [[halotolerance|tolerant to salinity]].<ref>{{cite journal|last=Rodriguez-Perez | first=J. A.|year=2009|title=Effect of Salinity on Seedling Emergence of Seven ''Banksia'' Species Cultivated for Cut Flower and Foliage|journal=Journal of Plant Nutrition|volume=32|issue=9|pages=1540–50|doi=10.1080/01904160903093844| s2cid=84912264}}</ref>
{{clear}}
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==External links==
{{Commons category
{{Wikispecies
* {{Wikisource-inline|Transactions of the Linnean Society of London/Volume 10/On the Proteaceae of Jussieu/Banksia#Banksia speciosa|Transactions of the Linnean Society of London/Volume 10/On the Proteaceae of Jussieu/Banksia#Banksia speciosa}}
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{{Taxonbar|from=Q4856722}}
[[Category:Banksia taxa by scientific name|speciosa]]
[[Category:Eudicots of Western Australia]]
[[Category:Trees of Australia]]
[[Category:Trees of Mediterranean climate]]
[[Category:Taxa named by Robert Brown (botanist, born 1773)]]
[[Category:Plants described in 1810]]
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