Banksia attenuata: Difference between revisions

| name = Candlestick Banksiabanksia or Biarabiara

| image = Banksia attenuata Marg River email.jpg

| image_caption = ''B. attenuata'',At [[Margaret River, Western Australia|Margaret River]]

|authority status_ref = [[Robert Brown (botanist, born 1773)|R.Br.]]<ref name="APC">{{cite webCite iucn|title=''Banksia attenuata'' |urlpage=https://biodiversitye.org.au/nsl/services/apc-format/display/106483T112520145A113306446|websitelast=Australian Plant CensusBarrett, S.|year=2020|access-date=2726 March 2022|doi=10.2305/IUCN.UK.2020-2.RLTS.T112520145A113306446.en}}</ref>

| range_map = B attentuata dist map.png

| range_map_caption = distributionDistribution of ''Banksia attenuata''

'''''Banksia attenuata''''', commonly known as the '''Candlestickcandlestick Banksiabanksia''', '''Slenderslender Banksiabanksia''', or '''Biarabiara''' to the [[NyoongarNoongar]] people,<ref isname=KP>{{cite aweb|url=https://www.bgpa.wa.gov.au/2014-08-29-05-50-09|title=Noongar glossary; Noongar Words and Definitions|accessdate=6 August 2022|publisher=[[speciesGovernment of Western Australia]]|year=2022|archive-date=6 August 2022|archive-url=https://web.archive.org/web/20220806122357/https://www.bgpa.wa.gov.au/2014-08-29-05-50-09|url-status=dead}}</ref> is a species of [[plant]] in the family [[Proteaceae]]. Commonly a [[tree]], it reaches {{convert|10|m|ft|abbr=on}} high, but it is often a [[shrub]] in drier areas {{convert|0.4|to|2|m|ft|abbr=on}} high. It has long, narrow, serrated [[leaf|leaves]] and bright yellow [[inflorescence]]s, or flower spikes, held above the foliage, which appear in spring and summer. The flower spikes age to grey and swell with the development of the woody [[follicle (fruit)|follicles]]. The candlestick banksia is found across much of the [[southwestSouthwest Australia|southwest]] of [[Western Australia]], from north of [[Kalbarri National Park]] down to [[Cape Leeuwin]] and across to [[Fitzgerald River National Park]].

The candlestick banksia is [[pollination|pollinated]] by and [[herbivore|provides food]] for a wide array of animals in summer months. Several species of [[honeyeater]] visit the flower spikes, as does the [[honey possum]], which has an important role as a pollinator. It regenerates from [[Bushfires in Australia|bushfire]] by regrowing from its woody base, known as a [[lignotuber]], or from [[epicormic bud]]s within its trunk. It can have a lifespan of 300 years. It has been widely used as a [[urban forestry|street tree]] and for [[amenity|amenities]] planting in urban Western Australia, though its large size generally precludes use in small gardens. A [[Dwarfing#Plants|dwarf form]] is commercially available in nurseries.

== Description ==

''Banksia attenuata'' is generally encountered as a tree up to {{convert|10|m|ft|abbr=on|sigfig=1}} tall. In the north of its range as the climate becomes warmer and drier, it is often a stunted multistemmed [[shrub]] {{convert|0.4|to|2|m|ft|abbr=on}} tall. Both forms occur in the vicinity of [[Hill River (Western Australia)|Hill River]] but there is otherwise a marked demarcation.<ref name="Cowl85">{{cite journal|last1=Cowling | first1=Richard M. | last2=Lamont | first2=Byron B. | author-link2=Byron Lamont |year=1985 |title=Variation in serotiny of three ''Banksia'' species along a climatic gradient |journal=Australian Journal of Ecology |volume=10 |issue=3 |pages=345–50 |doi=10.1111/j.1442-9993.1985.tb00895.x}}</ref>

[[File:Banksia attenuata tree.JPG|thumb|left|A large tree in [[Bold Park (Western Australia)|Bold Park]], Perth. The trunk is characteristically wavy or bent.|alt=A large tree with a wavy curved pale grey trunk in a dry scrubland type landscape]]

In the [[Wheatbelt (Western Australia)|Wheatbelt]] and east of the [[Stirling Range]], it is a stunted tree. Tree forms have a solid trunk, generally wavy or bent, with {{convert|1|–|2|cm|in|abbr=on|1}} thick crumbly orange-grey [[Bark (botany)|bark]] which is a red-brown underneath.<ref name="George81">{{The genus Banksia L.f. (Proteaceae)}}</ref> It regenerates from fire via [[lignotuber]] or epicormic buds from its fire-tolerant trunk. It has long narrow shiny green linear leaves {{convert|4|to|27|cm|in|abbr=on}} long and {{convert|0.5|to|1.6|cm|in|abbr=on|1}} wide.<ref name="George96" /><ref name=george99>{{cite encyclopedia| author=George, Alex S. |year=1999 |editor=Wilson, Annette |title=Flora of Australia | volume=17B |pages=175–251 |publisher=[[CSIRO Publishing]] / [[Australian Biological Resources Study]] |isbn=0-643-06454-0 |author-link=Alex George (botanist)|title-link=Flora of Australia (series) }}</ref> The leaf margins have v- or u-shaped serrations along their length. The new growth is a pale grey-green, and occurs mainly in the late spring and summer,<ref name=atlas>{{The Banksia Atlas}}, pp. 54–55</ref> often after [[flower]]ingflowering. The brilliant yellow [[inflorescence]]sinflorescences (flower spikes) occur from spring into summer and are up {{convert|5|cm|in|abbr=on}} wide and up to {{convert|25|–|30|cm|in|abbr=on}} tall.<ref name="George81" /> They are made up of many small individual flowers; a study at [[Mount Adams (Western Australia)|Mount Adams]] {{convert|330|km|mi|abbr=on}} north of [[Perth, Western Australia|Perth]] revealed a count of 1933 (± a [[Standard error (statistics)|standard error]] of 88) flowers per inflorescence,<ref name=Cowling87>{{cite journal|title= Seed bank dynamics in four co-occurring ''Banksia'' species|author1=Cowling, Richard M. |author2=Lamont, Byron B. |journal=Journal of Ecology |volume=75 |issue=2 |pages=289–302 |year=1987|jstor= 2260419|doi= 10.2307/2260419}} {{subscription required}}</ref> and another in the [[Fitzgerald River National Park]] yielded a count of 1720 (± 76) flowers. [[Anthesis]] proceeds up the flower spike over about 10 to 20 days, and is asynchronous. That is, a plant produces flower spikes over a several-week period and will thus have spikes at different stages of development over the flowering season.<ref name=Wooller01>{{cite journal|title= Seed set in two sympatric banksias, ''Banksia attenuata'' and ''B. baxteri'' |author1=Wooller, Sue J. |author2=Wooller, Ronald D | journal = Australian Journal of Botany |volume=49|issue=5|pages=597–602|year=2001|doi= 10.1071/BT00084}}</ref>

Often bright green in [[bud]] stage,<ref name=collins09>{{cite book |last1=Collins |first1=Kevin |last2=Collins |first2=Kathy |last3=George |first3=Alex S. |title=Banksias | publisher=Bloomings Books |location=Melbourne, Victoria |year=2008 |isbn=978-1-876473-68-6 |pages=68, 150–51}}</ref> they are terminal, occurring at the ends of one- to three-year-old branches, and displayed prominently above the foliage.<ref name="George81" /> The smell of the open flowers has been likened to a peppery [[Syrah|Shiraz]] wine.<ref name=collins09 /> Over time, the spikes fade to brown and then grey,<ref name="George96" /> and the individual flowers shrivel and lie against the spikes. This coincides with the development of dark furry oval [[follicle (fruit)|follicles]], which measure {{convert|2|–|3.5|cm|in|abbr=on|1}} long, {{convert|1|–|1.5|cm|in|abbr=on|1}} high, and {{convert|1.4|–|2|cm|in|abbr=on|1}} wide.<ref name="George81" /> However, only a very small percentage (0.1%) of flowers develop into follicles; the field study at Mount Adams yielded a count of 3.6 ± 1.2 per cone.<ref name=Cowling87 /> The follicles develop and mature over seven to eight months, from February to December, while [[seed]] development occurs over four months from September to December.<ref>{{cite journal|title=Seed developmental patterns in ''Banksia attenuata'' R.Br. and ''B. laricina'' C. Gardner in relation to mechanical defence costs |author1=Stock, W.D. |author2=Pate, J.S. |author3=Rasins, E. |journal=New Phytologist |volume=117 |issue=1 |pages=109–14 |year=1991 |doi=10.1111/j.1469-8137.1991.tb00950.x|doi-access=free }}</ref>

== Taxonomy ==

''Banksia attenuata'' was first collected by [[Robert Brown (botanist, born 1773)|Robert Brown]] from [[King George Sound (Western Australia)|King George Sound]] in December 1801, and published by him in 1810. The [[specific name (botany)|specific epithet]] is the [[Latin]] adjective ''attenuatus'' "'narrowed"', and refers to the leaves narrowing towards the base.<ref name="George96">{{cite book|last=George|first=Alex S.|title=The Banksia Book|edition=3rd|year=1996|publisher=Kangaroo Press|location=Kenthurst, New South Wales|isbn=0-86417-818-2|page=133}}</ref> The species has had a fairly uneventful [[taxonomy (biology)|taxonomic]] history. It has only two synonyms, and no [[subspecies]] or [[variety (botany)|varieties]] have been published;<ref name="APNI 52773">{{APNI | name = ''Banksia attenuata'' R.Br. | id = 52773}}</ref> Australian botanist [[Alex George (botanist)|Alex George]] reviewed the variation in form in the species, and felt that the tree and shrub forms differed only in size and hence were not distinct enough to represent separate [[taxon|taxa]].<ref name="George81" /> In 1840, [[John Lindley]] published a putative new species, ''Banksia cylindrostachya'', in his ''[[A Sketch of the Vegetation of the Swan River Colony]]''; this has now bebeen shown to be a [[taxonomic synonym]] of ''B.&nbsp;attenuata''.<ref name="APNI 53299">{{APNI | name = ''Banksia cylindrostachya'' Lindl. | id = 53299}}</ref> In 1891, [[Otto Kuntze]] rejected the generic name ''Banksia'' [[CarolusCarl Linnaeus the Younger|L.f.]], on the grounds that the name ''Banksia'' had previously been published in 1776 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, referring to this species as ''Sirmuellera attenuata''.<ref>{{cite book |last=Kuntze |first=Otto |authorlink=Otto Kuntze |title=Revisio generum plantarum |volume=2 |publisher=Arthur Felix |location=Leipzig |year=1891 |pages=581–582 |url=https://www.biodiversitylibrary.org/page/4004}}</ref> This application of the [[principle of priority]] was largely ignored by Kuntze's contemporaries,<ref>{{cite journal |last1=Rehder |first1=A. |authorlink1=Alfred Rehder |last2=Weatherby |first2=C. A. |authorlink2=Charles Alfred Weatherby |last3=Mansfeld |first3=R. |authorlink3=Rudolf Mansfeld |last4=Green |first4=M. L. |authorlink4=Mary Letitia Green |title=Conservation of Later Generic Homonyms |journal=Bulletin of Miscellaneous Information (Royal Botanic Gardens, Kew) |year=1935 |volume=1935 |issue=6/9 |page=368 |doi=10.2307/4107078 |jstor=4107078}}</ref> and ''Banksia'' L.f. was [[conserved name|formally conserved]] and ''Sirmuellera'' rejected in 1940.<ref>{{cite journal |last=Sprague |first=T. A. |authorlink=Thomas Archibald Sprague |title=Additional Nomina Generica Conservanda (Pteridophyta and Phanerogamae) |journal=Royal Botanic Gardens, Kew |year=1940 |volume=1940 |issue=3 |doi=10.2307/4111642 |jstor=4111642 |page=99}}</ref> [[Common name]]s include slender banksia, candle banksia, and candlestick banksia. ''Piara'' (alternately spelled ''biara'') is an [[Australian Aborigines|aboriginal]] name from the [[Melville, Western Australia|Melville]] region of Perth.

The relationships of ''Banksia attenuata'' within the genus are unclear. When [[Carl Meissner]] published [[Meissner's taxonomic arrangement of Banksia|his infrageneric arrangement]] of ''Banksia'' in 1856, he placed ''B.&nbsp;attenuata'' in section [[Banksia sect. Eubanksia|''Eubanksia'']] because its inflorescence is a spike rather than a domed head, and in series [[Banksia ser. Salicinae|''Salicinae'']],<ref name="Meissner 1856">{{cite encyclopedia |author=Meissner, Carl | author-link=Carl Meissner |year=1856 |chapter=Proteaceae |editor=de Candolle, A. P |encyclopedia=[[Prodromus systematis naturalis regni vegetabilis]] |volume=14 |location=Paris |publisher=Sumptibus Sociorum Treuttel et Wurtz |editor-link=A. P. de Candolle}}</ref> a large series that is now considered quite heterogeneous.<ref name="George81" /> This series was discarded in [[Bentham's taxonomic arrangement of Banksia|the 1870 arrangement]] of [[George Bentham]]; instead, ''B.&nbsp;attenuata'' was placed in section ''[[Banksia sectser. Cyrtostylis|''Cyrtostylis'']]'', a group of species whichthat did not fit easily into one of the other sections.<ref name="Bentham 1870">{{cite encyclopedia |last1=Bentham |first1=George | author-link=George Bentham |year=1870 |title=Banksia |encyclopedia=[[Flora Australiensis]] |volume=5 |pages=541–62 |location=London |publisher=L. Reeve & Co}}</ref>

George's arrangement remained current until 1996, when [[Kevin Thiele]] and [[Pauline Ladiges]] published an [[Thiele and Ladiges' taxonomic arrangement of Banksia|arrangement]] informed by a [[cladistics|cladistic]] analysis of [[morphology (biology)|morphological]] characteristics. They calculated ''B. attenuata'' to lie at the base of a large ''B. attenuata'' – ''B. ashbyi'' [[clade]], but conceded further work was needed before its relationships could be determined, and left it as ''[[incertae sedis]]'' (i.e. Its exact placement is unclear.).<ref name="Thiele 1996">{{cite journal |last1=Thiele |first1=Kevin | author-link1=Kevin Thiele |last2=Ladiges |first2=Pauline Y. | author-link2=Pauline Ladiges |year=1996 |title=A cladistic analysis of ''Banksia'' (Proteaceae) |journal=[[Australian Systematic Botany]] |volume=9 |issue=5 |pages=661–733 |doi=10.1071/SB9960661}}</ref> Questioning the emphasis on cladistics in Thiele and Ladiges' arrangement, George published a slightly modified version of his 1981 arrangement in his 1999 treatment of ''Banksia'' for the ''[[Flora of Australia (series)|Flora of Australia]]'' series of [[monograph]]s. To date, this remains the most recent comprehensive arrangement. The placement of ''B.&nbsp;attenuata'' in [[George's taxonomic arrangement of Banksia|George's 1999 arrangement]] may be summarised as follows:<ref name=george99 />

Since 1998, American botanist [[Austin Mast]] and co-authors have been publishing results of ongoing cladistic analyses of [[DNA sequence]] data for the subtribe [[Grevilleoideae|Banksiinae]], which then comprised genera ''Banksia'' and ''[[Banksia ser. Dryandra|Dryandra]]''. Their analyses suggest a [[phylogenetics|phylogeny]] that differs greatly from George's taxonomic arrangement. ''Banksia attenuata'' resolves as a [[basalBasal clade(phylogenetics)|basal]] member of and next closest relative, or 'sister', to a clade containing ''B.&nbsp;elegans'' and, within that, a [[monophyly|monophyletic]] ''B.''&nbsp;subg. ''Isostylis''.<ref name="Mast 1998">{{cite journal | last=Mast | first=Austin R. | author-link=Austin Mast |year=1998 |title=Molecular systematics of subtribe Banksiinae (''Banksia'' and ''Dryandra''; Proteaceae) based on cpDNA and nrDNA sequence data: implications for taxonomy and biogeography |journal=Australian Systematic Botany |volume=11 |issue=3–4 |pages=321–42 |doi =10.1071/SB97026}}</ref><ref name="Mast 2002">{{cite journal |last1=Mast | first1=Austin R. | last2=Givnish | first2=Thomas J. | author-link2=Thomas J. Givnish |year=2002 |title=Historical biogeography and the origin of stomatal distributions in ''Banksia'' and ''Dryandra'' (Proteaceae) based on their cpDNA phylogeny |journal=[[American Journal of Botany]] |volume=89 |issue=8 |pages=1311–23 |doi=10.3732/ajb.89.8.1311 |issn=0002-9122 |pmid=21665734|doi-access=free }}</ref><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 |publisher=CSIRO Publishing / Australian Systematic Botany Society |doi=10.1071/SB04015}}</ref> An [[Eocene]] fossil cone named ''[[Banksia archaeocarpa]]'', around 50 million years old, resembles that of ''B.&nbsp;attenuata''.<ref name="Wrigley 1991">{{cite book |last=Wrigley |first=John |author2=Fagg, Murray |title=Banksias, Waratahs and Grevilleas |year=1991 |publisher=Angus & Robertson |location=Sydney |isbn=0-207-17277-3 |pages=17, 88}}</ref>

Early in 2007, Mast and Thiele rearranged the genus ''Banksia'' by merging ''Dryandra'' into it, and published [[Banksia subg. Spathulatae|''B.''&nbsp;subg. ''Spathulatae'']] for the taxa having spoon-shaped cotyledons; thus ''B.''&nbsp;subg. ''Banksia'' was redefined as encompassing taxa lacking spoon-shaped cotyledons. They foreshadowed publishing a full arrangement once DNA sampling of ''Dryandra'' was complete;. inIn the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then ''B.&nbsp;attenuata'' is placed in ''B.''&nbsp;subg. ''Banksia''.<ref name="Mast 2007">{{cite journal |author1=Mast, Austin R. |author2=Thiele, Kevin |year=2007 |title=The transfer of ''Dryandra'' R.Br. to ''Banksia'' L.f. (Proteaceae) |journal = Australian Systematic Botany |volume=20 |issue=1 |pages=63–71 |doi=10.1071/SB06016}}</ref>

== Distribution and habitat ==

== Ecology ==

Like many plants in [[south-westsouthwest Western Australia]], ''B.&nbsp;attenuata'' is adapted to an environment in which bushfire events are relatively frequent. Most ''Banksia'' species can be placed in one of two broad groups according to their response to fire: ''reseeders'' are killed by fire, but fire also triggers the release of their [[canopy seed bank]], thus promoting recruitment of the next generation; ''[[resprouter]]s'' survive fire, resprouting from a [[lignotuber]] or, more rarely, [[Epicormic shoot|epicormic buds]] protected by thick bark.<ref name="Lamont 1995">{{cite journal |last1=Lamont |firstfirst1=Byron B. |last2=Markey |first2=Adrienne |year=1995 |title=Biogeography of fire-killed and resprouting ''Banksia'' species in South-western Australia |journal=Australian Journal of Botany |volume=43 |issue=3 |pages=283–303 |doi=10.1071/BT9950283}}</ref> Bearing epicormic buds and a lignotuber, ''B.&nbsp;attenuata'' is one of the latter group, with follicles that may open spontaneously or by fire.<ref name="George81" />

Like many members of the family Proteaceae, ''Banksia attenuata'' is an obligate [[outcrossing]] species. [[Self-incompatibility in plants|Self-incompatible]], inflorescences require pollinators to be fertilised and produce seed. A genetic study of seed collected near [[Jandakot, Western Australia|Jandakot]] published in 1980 showed obligate outcrossing.<ref>{{cite journal|title= Estimation of the outcrossing oate for ''Banksia attenuata'' R. Br. and ''Banksia menziesii'' R. Br. (Proteaceae)|author=Scott, John K.|journal= Australian Journal of Botany |volume=28|issue=1|pages=53–59|year=1980|doi= 10.1071/BT9800053}}</ref> A field study in the Fitzgerald River National Park where inflorescences were enclosed in mesh fine enough to keep out vertebrates and invertebrates as small as the honey bee (''Apis mellifera'') showed that follicles still developed, indicating that small invertebrates were able to cross-pollinate the species.<ref name=Wooller01 /> ''Banksia attenuata'' flowers are visited by the [[colletidae|colletid]] bee ''[[Hylaeus globuliferus]]'' and bees of the genus ''[[Euhesma]]''.<ref>{{cite web|url=http://flyaqis.museum.vic.gov.au/cgi-bin/texhtml|title=Specimen Report|year=2010|work=Museum Victoria website: Bioinformatics|publisher=Museum Victoria|access-date=25 November 2010|location=Melbourne, Australia|archive-date=15 June 2009|archive-url=http://webarchive.loc.gov/all/20090615224236/http://flyaqis.museum.vic.gov.au/cgi-bin/texhtml|url-status=dead}}</ref> Other pollinating invertebrates recorded include ants and dragonflies.<ref name=atlas /> An analysis of the invertebrate population found in the canopy of ''Banksia'' woodland found that mites and ticks ([[acari]]), beetles ([[coleoptera]]) and ants, bees and wasps ([[hymenoptera]]) predominated overall, with the three orders also the most common on ''B. attenuata'' along with thrips ([[thysanoptera]]). Lower overall numbers of invertebrates on ''Banksia'' species were thought to be related to the presence of insectivorous birds.<ref>{{cite journal|title=Abundance of Arthropods in Tree Canopies of ''Banksia'' Woodland on the Swan Coastal Plain|author1=Tassone, R.A. |author2=Maier, J.D. |journal=Journal of the Royal Society of Western Australia |volume=80 |pages=281–86 |year=1997}}</ref>

Many bird species were recorded by the national Banksia Atlas survey, including the [[New Holland honeyeater]] (''Phylidonyris novaehollandiae''), [[brown honeyeater]] (''Lichmera indistincta''), [[singing honeyeater]] (''LichenostomusGavicalis virescens''), [[western spinebill]] (''Acanthorhynchus superciliosus''), [[Australian ringneck|twenty-eight parrot]] (''Barnardius zonarius semitorquatus'') and [[red-tailed black cockatoo]] (''Calyptorhynchus banksii'').<ref name=atlas /> Black cockatoos have been observed feeding upon the seed of ''B.&nbsp;attenuata'', although it is not clear which species of black cockatoo was observed, the short-billed or [[long-billed black cockatoo]] (''Calyptorhynchus baudinii'').<ref name="Barker 1984">{{cite book |author1=Barker, R.D. |author2=Vestjens, W.J.M. | year = 1984 | title = The Food of Australian Birds | publisher = Melbourne University Press | isbn = 0-643-05006-X | pages = '''1''':331, '''2''':458}}</ref> At a site near Jandakot, short-billed black cockatoos were observed selecting immature infructescences which bore signs of infestation by the weevil ''[[Alphitopis nivea]]'', the larvae of which tunnel in banksia spikes and eat the seed. They extract the larvae and drop the cones.<ref>{{cite journal|title=Selective predation by White-tailed Black Cockatoos on fruit of ''Banksia attenuata'' containing the seed-eating weevil ''Alphitopis nivea'' |author1=Scott, John K. |author2=Black, Robert | journal=Australian Wildlife Research| volume=8 |issue=2 |pages=421–30 |year=1981 |doi=10.1071/WR9810421}}</ref>

A 1978 field study conducted around [[Albany, Western Australia|Albany]] found the [[honey possum]] (''Tarsipes rostratus'') was a major pollinator of ''Banksia attenuata'', both feeding directly on the pollen and drinking the nectar. The flower structure is suited to passing pollen onto the possum as it feeds, unlike honeyeaters whose bills are too long for this to occur readily. Coupled with the flower spike's musky odour, these findings suggest ''Banksia attenuata'' is highly adapted to be pollinated primarily by this mammal species.<ref name=Weins79>{{cite journal|author1=Weins, Delbert |author2=Renfree, Marilyn |author3=Wooller, Ronald D. |year=1979 |title=Pollen loads of Honey possums (''Tarsipes spencerae'') and non-flying mammal pollination in South-western Australia |journal=Annals of the Missouri Botanical Garden|volume=66|pages=830–38 |url=http://www.botanicus.org/item/31753003566442 |access-date=24 November 2010|doi=10.2307/2398921|issue=4|jstor=2398921 }}</ref> Furthermore, Petroc Sumner and colleagues have investigated the [[Cone cell|cone]] [[photoreceptor cell]]s of honey possums and compared them with the colour changes of ''B.&nbsp;attenuata''. They found that the possum is trichromatic (like humans and possibly many marsupials) and propose that its L (long wavelength) cones help it discern ''B.&nbsp;attenuata'' flower spikes, and the M (or medium wavelength) cones could help it distinguish ripe inflorescences with nectar from recently finished spikes (a difficult task for human vision).<ref>{{cite journal|title=The ecology of visual pigment tuning in an Australian marsupial: the honey possum ''Tarsipes rostratus'' |author1=Sumner, Petroc |author2=Arrese, Catherine A. |author3=Partridge, Julian C. |journal = Journal of Experimental Biology|volume=208| issue=10|pages=1803–15|year=2005|doi=10.1242/jeb.01610|url=http://jeb.biologists.org/cgi/content/full/208/10/1803 | format=fulltext |access-date=4 December 2010 |pmid=15879062}}</ref> There is some evidence for other mammals as pollinators; ''B.&nbsp;attenuata''-like pollen was recovered from museum skins of [[dunnart]]s (''Sminthopsis'' spp.) and pygmy possums (''[[Cercartetus]]'' spp.), and the [[sugar glider]] (''Petaurus breviceps'') visited flower spikes in captivity.<ref name=Weins79 />

All banksias have developed proteoid or cluster roots in response to the nutrient-poor conditions of Australian soils (particularly lacking in [[phosphorus]]).<ref>{{cite journal|last=Beadle, Noel C.|year=1968|title=Some aspects of the ecology and physiology of Australian xeromorphic plants |journal=Australian Journal of Science|volume=30|pages=348–355}}</ref> These have been measured in Eneabba extending to a depth of {{convert|15|cm|in|abbr=on}} below the soil surface.<ref>{{cite journal|title=Aerial and below-ground phytomass of ''Banksia'' Scrub-heath at Eneabba, South-western Australia|author1=Low, A.B. |author2=Lamont, Byron B. |journal=Australian Journal of Botany |volume=38 |issue=4 |pages=351–59 |year=1990|doi=10.1071/BT9900351}}</ref> The plant develops masses of fine lateral roots which form a mat-like structure underneath the soil surface, and enable it to extract nutrients as efficiently possible out of the soil. A study of three co-occurring species in ''Banksia'' woodland in southwestern Australia—''Banksia menziesii'', ''B.&nbsp;attenuata'' and ''B.&nbsp;ilicifolia''—found that all three develop fresh roots in September after winter rainfall, and that the bacteria populations associated with the root systems of ''B.&nbsp;menziesii'' differ from the other two, and that they also change depending on the age of the roots.<ref>{{cite journal|author1=Marschner, Petra |author2=Grierson, Pauline F. |author3=Rengel, Zed |year=2006|title=Microbial community composition and functioning in the rhizosphere of three ''Banksia'' species in native woodland in Western Australia |journal=Applied Soil Ecology|volume=28|issue=3|pages=191–201|doi=10.1016/j.apsoil.2004.09.001}}</ref> Another study on root architecture of ''Banksia hookeriana'', ''B.&nbsp;menziesii'' and ''B.&nbsp;attenuata'' found the overall structure of all three to be similar, with proteoid mats more active and growing in wetter months (winter-spring). Plants send out several sinker roots which descend to reach the [[water table]], and the original tap root may or may not have died off.<ref>{{cite journal|author1=Lamont, Byron B. |author2=Bergl, Stephen M. |year=1991 |title=Water relations, shoot and root architecture, and phenology of three co-occurring ''Banksia'' species: no evidence for niche differentiation in the pattern of water use |journal=Oikos |volume=60 |issue=3 |pages=291–98 |jstor=3545070|doi=10.2307/3545070}} {{subscription required}}</ref> Along with ''B.&nbsp;menziesii'', ''B.&nbsp;attenuata'' is a facultative [[phreatophyte]]. The two species are less strictly tied to the [[water table]] and hence able to grow in a wider variety of places within ''Banksia'' woodland habitat around Perth than the co-occurring ''[[Banksia ilicifolia|B.&nbsp;ilicifolia]]'' and ''[[Banksia littoralis|B.&nbsp;littoralis]]''.<ref>{{cite journal |author1=Froend, R.H. |author2=Drake, P.L. |year=2006 |title=Defining phreatophyte response to reduced water availability: preliminary investigations on the use of xylem cavitation vulnerability in ''Banksia'' woodland species |journal=Australian Journal of Botany |volume=54 |issue=2 |pages=173–79 |doi=10.1071/BT05081}}</ref> A study at a rehabilitation site on a sand mine north of Perth found that the broadleaved species ''B.&nbsp;attenuata'' and ''B.&nbsp;hookeriana'' were harder to establish than the fine-leaved ''[[Banksia leptophylla|B.&nbsp;leptophylla]]'' there, due to increased impedance of the disturbed soil.<ref>{{cite journal|title=Survival, growth and water relations of ''Banksia'' seedlings on a sand mine rehabilitation site and adjacent scrub-heath sites |author1=Enright, N.J. |author2=Lamont, Byron B. | journal=Journal of Applied Ecology |volume=29 |issue=3 |pages=663–71 |year=1992 |jstor=2404474|doi=10.2307/2404474}} {{subscription required}}</ref> Analysis of native plant species at a remnant of banksia woodland in suburban Perth which had been invaded by two herbaceous weed species (''[[Ehrharta calycina]]'' and ''[[Pelargonium capitatum]]'') found increased [[phosphorus]] levels in native foliage. Although ''B.&nbsp;attenuata'' leaves did not have increased phosphorus, they did have reduced levels of [[manganese]]—an element which is absorbed into the plant by its proteoid roots, the formation of which can be inhibited by raised levels of phosphorus.<ref>{{cite journal |title=Enhanced soil and leaf nutrient status of a Western Australian Banksia woodland community invaded by'' Ehrharta calycina'' and ''Pelargonium capitatum''|author1=Fisher, Judith L. |author2=Veneklaas Erik J. |author3=Lambers, Hans |author4=Loneragan, William A. |journal=Plant and Soil |volume=284 |issue=1–2 |pages=253–64 |year=2006 | format=fulltext |access-date=1 December 2010 |doi=10.1007/s11104-006-0042-z}}</ref>

In a 1985 study inoculating cultivated plants, ''Banksia attenuata'' showed moderate to high susceptibility to ''[[Phytophthora cinnamomi]]'' dieback;<ref name="McCredie 1985">{{cite journal |last1=McCredie |first1=T.A. |last2=Dixon |first2=K.W. |last3=Sivasithamparam |first3=K. |year=1985 |title=Variability in the resistance of ''Banksia'' L.f. species to ''Phytophthora cinnamomi'' Rands |journal=[[Australian Journal of Botany]] |volume=33 |issue=6 |pages=629–37 |doi=10.1071/BT9850629}}</ref> and at least some field and cultivation evidence points to it being highly susceptible.<ref name="DEH 2006">{{cite web |year=2006 |title=Part 2, Appendix 4: The responses of native Australian plant species to ''Phytophthora cinnamomi'' |work=Management of Phytophthora cinnamomi for Biodiversity Conservation in Australia |publisher=[[Department of the Environment and Heritage]], Australian Government |url=http://www.environment.gov.au/biodiversity/invasive/publications/pubs/appendix4.pdf |format=PDF |access-date=22 August 2009}}</ref> ''P.&nbsp;cinnamomi'' spreads from plant to plant via lateral roots, advancing at a rate of around a metre a year. The symptoms of infection in ''Banksia attenuata'' include yellowing of the leaves in the tree crown, and lesions at the base of the trunk. The red healthy roots become discoloured brown.<ref>{{cite journal|title=Invasion of Bassendean dune Banksia woodland by ''Phytophthora cinnamomi'' |author1=Hill, T.C.J. |author2=Tippett, J.T. |author3=Shearer, B.L. |journal=Australian Journal of Botany |volume=42 |issue=6 |pages=725–38 |year=1994 |doi=10.1071/BT9940725}}</ref> A study of ''Banksia attenuata'' woodland {{convert|400|km|mi|abbr=on}} southeast of Perth across 16 years and following a wave of ''P.&nbsp;cinnamomi'' infestation showed that ''B.&nbsp;attenuata'' populations still existed but were significantly reduced in diseased areas.<ref>{{cite journal|title=Community-level changes in Banksia woodland following plant pathogen invasion in the Southwest Australian Floristic Region|author= Bishop, C.L. |author2=Wardell-Johnson, G.W. |author3=Williams, M.R. |journal=Journal of Vegetation Science|volume=21|issue=5|pages=888–98| year=2010|doi=10.1111/j.1654-1103.2010.01194.x}}</ref> Injecting a solution of [[phosphite]] into the trunks of affected ''B.&nbsp;attenuata'' trees at a disease front in ''Banksia'' woodland can delay morbidity from dieback for five years.<ref>{{cite journal|title=A stem injection of phosphite protects ''Banksia'' species and ''Eucalyptus marginata'' from ''Phytophthora cinnamomi'' for at least four years|author1=Shearer, Bryan L. |author2=Fairman, Richard G. |journal=Australasian Plant Pathology |volume=36 |issue=1 |pages=78–86 |year=2007 |doi=10.1071/AP06085}}</ref> Injecting and spraying phosphite also reduces the rate of spread of a dieback front for around five years. A bushfire did not influence this slowing.<ref>{{cite journal |title=Phosphite reduces disease extension of a ''Phytophthora cinnamomi'' front in Banksia woodland, even after fire |author1=Shearer, Bryan L. |author2=Fairman, Richard G. |journal=Australasian Plant Pathology |volume=33 |issue=2 |pages=249–54 |year=2004 |doi=10.1071/AP04002}}</ref> A 2003 study found that drenching the soil with 0.50&nbsp;[[Molar concentration#Units|mM]] [[benzoic acid]] significantly reduced the size of ''P.&nbsp;cinnamomi'' lesions.<ref>{{cite journal|title=Benzoic acid induces tolerance to biotic stress caused by ''Phytophthora cinnamomi'' in ''Banksia attenuata''|author1=Williams, Mia |author2=Senaratna, Tissa |author3=Dixon, Kingsley |author4=Sivasithamparam, Krishnapillai |journal=Plant Growth Regulation |volume=41 |issue=1 |pages=89–91 |doi=10.1023/A:1027355604096

The well-displayed bright yellow spikes are an attractive feature, with shrubby dwarf forms more versatile horticulturally.<ref name=matth>{{cite book |author=Matthews, Lewis J. |title=The Protea Book: A Guide to Cultivated Proteaceae |page=39 |isbn=0-88192-553-5 |year=2002 |publisher=Timber Press |location=Portland, Oregon |url=https://books.google.com/books?id=vGBV3kkKcIcC&pg=PA39&dq=%22Banksia+attenuata%22&pg=PA39}}</ref> All forms of ''Banksia attenuata'' require good drainage, sandy soil and a sunny position to do well, with a [[pH]] between 5.5 and 7.0.<ref name=collins09 /> They are sensitive to [[Phytophthora cinnamomi|dieback]], and do not fare well in humid climates.<ref name="Wrigley 1991" /> Seeds do not require any treatment, and take 16 to 49 days to [[germinate]].<ref name="Sweedman 2006">{{cite book |author1=Sweedman, Luke |author2=Merritt, David |year=2006 |title=Australian Seeds: A Guide to their Collection, Identification and Biology |publisher=CSIRO Publishing |isbn=0-643-09298-6 |page=202}}</ref> Seedlings are highly vulnerable to [[damping off]].<ref name="Wrigley 1991" /> Plants take from four to six years to flower from seed.<ref name=collins09 /> There has been little success with other methods of improving adaptability to humid climates such as [[grafting]].<ref name="Wrigley 1991" /> Flower spikes in late bud are used in the [[cut flower]] industry,<ref name=collins09 /> primarily in Western Australia.<ref name=matth />

== External links ==

{{WikisourceparWikisource|Transactions of the Linnean Society of London/Volume 10/On the Proteaceae of Jussieu/Banksia#Banksia_attenuata|Transactions of the Linnean Society of London/ Volume 10/ On the Proteaceae of Jussieu/ Banksia#Banksia_attenuata}}

{{WikisourceparWikisource|Edwards's Botanical Register/Appendix to the First Twenty-three Volumes/A Sketch of the Vegetation of the Swan River Colony/Proteaceae#Banksia cylindrastachya| A Sketch of the Vegetation of the Swan River Colony/ Proteaceae#Banksia cylindrastachya}}

{{WikisourceparWikisource|Flora Australiensis/Volume 5/Proteaceae/Banksia#Banksia attenuata|Flora Australiensis/ Volume 5/Proteaceae/ Banksia#Banksia attenuata}}

{{Wikispecies-inline|Banksia attenuata}}

{{Commons category-inline}}

* {{Flora of Australia Online|name=''Banksia attenuata'' R.Br.}}

* {{FloraBase|name=''Banksia attenuata'' R.Br.}}