Taxonomy of Banksia
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[[Image:Banksia serrata watercolour from Bank's Florilegium.jpg|thumb|right|A 1770s watercolour of ''[[Banksia serrata]]'', the [[type species]] of ''[[Banksia]]''. One of the first ever drawings of a ''Banksia'' species, it was produced by one of Sir [[Joseph Banks]]' London artists, based on original drawings by [[Sydney Parkinson]], who was present when the genus was first collected at [[Botany Bay]].]]
As with other [[flowering plant]]s, the '''taxonomy of ''Banksia''''' is based on [[anatomy|anatomical]] and [[comparative anatomy|morphological]] properties of the ''[[Banksia]]'' [[flower]], [[fruit|fruiting structure]] and [[seed]], along with secondary characteristics such as leaf structure and growth habit. The genus is placed in family [[Proteaceae]], subfamily [[Grevilleoideae]], tribe [[Banksieae]] and subtribe [[Banksiinae]], alongside its close relative ''[[Dryandra]]''. The currently accepted infrageneric arrangement is based on [[Alex George]]'s 1999 monograph for the ''[[Flora of Australia (series)|Flora of Australia]]'' book series, and recognises two subgenera, three sections, 13 series, 77 species, 6 subspecies and 18 varieties. Recent [[cladistics|cladistic analyses]] have cast doubt on aspects of this arrangement, but proposed alternative arrangements have not been widely accepted.
==Background==
{{seealso|Banksia}}
''Banksia'' is a [[genus]] of around 80 [[species]] in the [[plant]] family [[Proteaceae]]. An iconic [[Australia]]n wildflower and popular [[garden]] [[plant]], they are easily recognised by their characteristic flower spikes and fruiting "cones". They grow in forms varying from prostrate woody [[shrub]]s to trees up to 35 metres tall, and occur in all but the most [[arid]] areas of Australia. As heavy producers of [[nectar (plant)|nectar]], they are important sources of food for nectariferous animals such as [[honeyeater]]s and [[honey possum]]s, and they are of economic importance to the [[nursery (horticulture)|nursery]] and [[floristry|cut flower]] industries. However they are threatened by a number of processes, including land clearing, frequent burning, and disease; and a number of species are [[rare species|rare]] and [[endangered species|endangered]].
==Taxonomic history==
{{see also|Timeline of Banksia}}
[[Image:Supplementum Plantarum Systematis Vegetabilium.jpg|thumb|right|''Banksia'' was first described in [[Carolus Linnaeus the Younger]]'s April 1782 publication ''[[Supplementum Plantarum]]''.]]
Specimens of ''Banksia'' were first collected by [[Joseph Banks|Sir Joseph Banks]] and [[Daniel Solander|Dr Daniel Solander]], naturalists on the ''[[HM Bark Endeavour|Endeavour]]'' during Lieutenant (later Captain) [[James Cook]]'s first voyage to the [[Pacific Ocean]]. Cook landed on Australian soil for the first time on [[29 April]] [[1770]], at a place that he later named [[Botany Bay]] in recognition of "the great quantity of plants Mr Banks and Dr Solander found in this place".<ref name="Wharton 1893">{{cite book | author = [[James Cook|Cook, James]] | year = 1893 | editor = William J. L. Wharton (ed.) | title = [[Wikisource:Captain Cook's Journal, First Voyage|Captain Cook's Journal during his First Voyage Round the World made in H. M. Bark "Endeavour" 1768-71: A Literal Transcription of the Original MSS]] | location = London | publisher = E. Stock}}</ref> Over the next seven weeks, Banks and Solander collected thousands of plant specimens, including the first specimens of a new genus that would later be named ''Banksia'' in Banks' honour. Four species were present in this first collection: ''[[Banksia serrata|B. serrata]]'' (Saw Banksia), ''[[Banksia integrifolia|B. integrifolia]]'' (Coast Banksia), ''[[Banksia ericifolia|B. ericifolia]]'' (Heath-leaved Banksia) and ''[[Banksia robur|B. robur]]'' (Swamp Banksia). In June the ship was [[careening|careened]] at [[Endeavour River]], where specimens of ''[[Banksia dentata|B. dentata]]'' (Tropical Banksia) were collected.
Every specimen collected during the ''Endeavour'' voyage was sketched by Banks' [[botanical illustrator]] [[Sydney Parkinson]]. On the ''Endeavour''<span/>'s return to [[England]] in July 1771, Banks' specimens became part of his [[London]] [[herbarium]], and artists were employed to paint [[watercolour]]s from Parkinson's sketches. Banks had plans to publish his entire collection as "[[Banks' Florilegium]]", but for various reasons the project was never completed, and it would be ten years before any of the ''Banksia'' species were formally published.<ref name="Salkin 1981">{{cite journal | author = Salkin, A. I. | year = 1981 | title = A Short History of the Discovery and Naming of Banksias in Eastern Australia: Part I, Banks & Solander | journal = Victorian Naturalist | volume = 98 | issue = 2}}</ref> By this time, a sixth species had been collected; in 1776, during Cook's third voyage, [[David Nelson]] collected specimens of ''[[Banksia marginata|B. marginata]]'' (Silver Banksia) from [[South Bruny Island]], [[Tasmania]].
The ''Banksia'' genus was finally described and named by [[Carolus Linnaeus the Younger]] in his April 1782 publication ''[[Supplementum Plantarum]]''; hence the [[Binomial nomenclature#Authorship in scientific names|full name]] for the genus is "''Banksia'' L.f.". Linnaeus placed the genus in class Tetrandra, order Monogynia of his father's classification,<ref name="L.f. 1782">{{cite book | author = [[Carolus Linnaeus the Younger]] | year = 1782 | title = [[Supplementum Plantarum|Supplementum Plantarum Systema Vegetabilium Editionis Decima Tertia, Generum Plantarum Editionis Fexta, Et Specierum Plantarum Editionis Secunda]] | location = Brunsvigae |publisher = Orphanotrophei}}</ref> and named it in honour of Banks. The name ''Banksia'' had in fact already been published in 1775 as ''Banksia'' [[Johann Reinhold Forster|J.R.Forst]] & [[Georg Forster|G.Forst]], referring to some [[New Zealand]] species that the Forsters had collected during Cook's second voyage. However Linnaeus incorrectly attributed the Forsters' specimens to the genus ''[[Passerina (plant)|Passerina]]'', and therefore considered the name ''Banksia'' available for use. By the time [[Joseph Gaertner]] corrected Banks' error in 1788, ''Banksia'' L.f. was widely known and accepted, so Gaertner [[nomen novum|renamed]] ''Banksia'' J.R.Forst & G.Forst to ''[[Pimelia]]'', a name previously chosen for the genus by Banks and Solander.<ref name="Salkin 1981">{{cite journal | author = Salkin, A. I. | year = 1981 | title = A Short History of the Discovery and Naming of Banksias in Eastern Australia: Part I, Banks & Solander | journal = Victorian Naturalist | volume = 98 | issue = 2}}</ref>
''Banksia'' L.f. has since been challenged a number of times. The later near-homonym ''Banksea'' Koenig was published in 1783, but subsequently determined to be a synonym of ''[[Costus]]'' L. In 1790 [[James Bruce]] published ''Bankesia'' Bruce, later corrected to ''Banksia'' Bruce, but the name was rejected in favour of [[Johann Friedrich Gmelin]]'s name ''[[Hagenia]]''. In 1820 the name ''Banksia'' [[Joseph Dombey|Dombey]] ex [[A. P. de Candolle|DC.]] was published, but this was later determined by be a [[nomen nudum]] that referred to the genus ''[[Cuphea]]'', In 1891, [[Otto Kuntze]] proposed to enforce the right of precedent of ''Banksia'' J.R.Forst & G.Forst, renaming ''Pimelea'' to ''Banksia'', and proposing the name ''Sirmuellera'' Kuntze in place of ''Banksia'' L.f. This challenge failed, as did [[James Britten]]'s 1905 challenge.<ref name="George 1981B">{{cite book | author = George, A. S. | year = 1981 | chapter = The genus ''Banksia'' L.f. — a case history in Australian botany | title = History in the service of systematics : papers from the Conference to celebrate the centenary of the British Museum (Natural History | pages = 53-59| isbn= 0-901843-05-9| location=London| publisher=Society for the Bibliography of Natural History}}</ref> In 1940, ''Banksia'' L.f. was formally conserved against ''Banksia'' J.R.Forst. & G.Forst by [[Thomas Sprague]].<ref name="Olde 2002"/>
In 1810, [[Robert Brown (botanist)|Robert Brown]] published descriptions and [[Brown's taxonomic arrangement of Banksia|a taxonomic arrangement]] of the 31 known species of ''Banksia'' in his ''[[Prodromus Florae Novae Hollandiae et Insulae Van Diemen]]''. He placed ''[[Banksia ilicifolia|B. ilicifolia]]'' alone in subgenus [[Banksia subg. Isostylis|''Isostylis'']], in recognition of its unusual dome-shaped [[inflorescence]]. All other species were placed in subgenus ''Banksia verae'', the "True Banksias". Brown made no attempt to classify the species below the subgenus level. He described another eleven ''Banksia'' species in his 1830 supplement, placing them all in ''Banksia verae'' in accordance with his 1810 classification.<ref name="Thiele 1996"/>. ''Banksia verae'' was renamed ''Eubanksia'' by [[Stephan Ladislaus Endlicher|Stephan Endlicher]] in 1847.
By the time [[Carl Meissner]] published his 1856 classification of the Proteaceae, there were 58 described ''Banksia'' species. [[Meissner's taxonomic arrangement of Banksia|Meissner's arrangement]] gave ''Isostylis'' and ''Eubanksia'' sectional rank, and divided the latter into four series based on leaf properties; these series were all highly heterogeneous.<ref name="Thiele 1996"/>
[[George Bentham]] published [[Bentham's taxonomic arrangement of Banksia|his arrangement]] of the ''Banksia'' in his landmark 1870 publication ''[[Flora Australiensis]]''.<ref name="Bentham 1870">{{cite encyclopedia | first = George|last= Bentham|authorlink= George Bentham| year = 1870 | title = Banksia| encyclopedia= [[Flora Australiensis|Flora Australiensis: A Description of the Plants of the Australian Territory]] | volume = Volume 5: Myoporineae to Proteaceae | pages=541–562 | location = London | publisher = L. Reeve & Co.}}</ref> The number of recognised ''Banksia'' species was reduced to 46, and Meissner's four heterogeneous series were replaced by four sections based on leaf, style and [[pollen-presenter]] characters, taking the number of sections to five. Three of these sections were fairly well-defined and homogeneous, while another, ''Orthostylis'', was somewhat heterogeneous. The fourth, ''Cyrtostylis'', was erected to contain the species that did not belong in the other sections, and was therefore highly heterogeneous.<ref name="Thiele 1996"/> Despite these shortcomings, this arrangement would stand for over 100 years.
==Classification and relationships within Proteaceae==
[[Image:Dryandra sessilis 2 cape Naturaliste email.jpg|thumb|right|''Banksia'' is closely related to ''[[Dryandra]]'' (''[[Dryandra sessilis|D. sessilis]]'' pictured). In fact the two genera are so closely related that there have been calls for ''Dryandra'' to be merged into ''Banksia''.]]
The framework for classification of genera within [[Proteaceae]] was laid in 1975 by [[Lawrence Alexander Sidney Johnson|L. A. S. Johnson]] and [[Barbara G. Briggs|Barbara Briggs]].<ref name="Lawrence 1975">{{cite journal|first = L. A. S. | last = Johnson | authorlink = Lawrence Alexander Sidney Johnson | coauthors = [[Barbara G. Briggs|Barbara Briggs]] | year = 1975 | title = On the Proteaceae: The evolution and classification of a southern family | journal= Journal of the Linnean Society of London. Botany | volume = 70 | pages = 83–182|id={{ISSN|0368-2927}} | doi = 10.1111/j.1095-8339.1975.tb01644.x}}</ref><!--Issue number is missing--> Their classification has been refined somewhat over the ensuing three decades, resulting in a fairly stable and widely accepted arrangement. Proteaceae is divided into seven subfamilies, with ''Banksia'' placed in subfamily [[Grevilleoideae]] because the individual flowers in its inflorescence occur in pairs. On the basis of certain characters of the [[leaf#Veins|leaf venation]], hairs and [[pollen]], it is grouped with three other genera in the tribe [[Banksieae]]. Two small genera, ''[[Austromuellera]]'' and ''[[Musgravea]]'', both of which occur only in the [[rainforest]]s of [[Queensland]], are placed in subtribe [[Musgraveinae]]. ''Banksia'' and ''[[Dryandra]]'' are placed in subtribe [[Banksiinae]] on a number of grounds of which the most obvious and easily recognised is the occurrence of flowers in condensed heads.<ref name="George 1999"/> The placement of ''Banksia'' in Proteaceae can be summarised as follows:
:'''Family [[Proteaceae]]'''
::Subfamily [[Persoonioideae]]
::Subfamily [[Bellendenoideae]]
::Subfamily [[Eidotheoideae]]
::Subfamily [[Proteoideae]]
::Subfamily [[Sphalmioideae]]
::Subfamily [[Carnarvonioideae]]
::'''Subfamily [[Grevilleoideae]]'''
:::Tribe [[Oriteae]]
:::Tribe [[Knightieae]]
:::Tribe [[Embothrieae]]
:::Tribe [[Helicieae]]
:::Tribe [[Macadamieae]]
:::Tribe [[Grevilleeae]]
:::'''Tribe [[Banksieae]]'''
::::Subtribe [[Musgraveinae]]
:::::Genus ''[[Austromuellera]]''
:::::Genus ''[[Musgravea]]''
::::'''Subtribe [[Banksiinae]]'''
:::::'''Genus ''[[Banksia]]'''''
:::::Genus ''[[Dryandra]]''
Although the taxonomic legitimacy of tribe Banksiinae is universally recognised, there has been some debate about the legitimacy of the tribe's resolution into genera ''Banksia'' and ''Dryandra''. For a number of years this debate centred on similarities between the inflorescences of ''Banksia'' subg. ''Isostylis'' species and those of ''Dryandra''. These similarities led to calls for the genera to be merged, or for ''Isostylis'' to be moved across to ''Dryandra''. However, Alex George and other supporters of the ''status quo'' argued that the similarities between ''Isostylis'' and ''Dryandra'' were matters of superficial appearance, whereas similarities between ''Isostylis'' and other ''Banksia'' species were far more important diagnostically.<ref name="George 1981">{{The genus Banksia L.f. (Proteaceae)}}</ref> Recent [[DNA]] analyses led by Austin Mast have confirmed George's position that ''Dryandra'' and ''Isostylis'' are not especially closely related, but have also provided powerful evidence that ''Banksia'' is paraphyletic with ''Dryandra''. Mast has advised that the least disruptive approach to restore [[monophyly]] would be to sink ''Dryandra'' into ''Banksia''.<ref name="Mast 2002">{{cite journal | first = Austin R. | last = Mast | coauthors = Thomas J. Givnish | month = August| year = 2002 | title = Historical Biogeography and the Origin of Stomatal Distributions in Banksia & Dryandra (Proteaceae) Based on Their cpDNA Phylogeny | journal = American Journal of Botany | volume = 89 | issue = 8 | pages = 1311–1323 | id = {{ISSN|0002-9122}} | url = http://www.amjbot.org/cgi/content/full/89/8/1311|accessdate=2006-07-02 | doi = 10.3732/ajb.89.8.1311}}</ref><ref name="Mast 2005">{{cite journal | author = [[Austin Mast|Mast, Austin R.]], Eric H. Jones and Shawn P. Havery | 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 | id={{ISSN|1030-1887}} | doi = 10.1071/SB04015}}</ref>
==Classical taxonomic treatment==
[[Image:B ericifolia erowal bay1.JPG|thumb|right|''B. ericifolia'' is placed in section ''Oncostylis'' on the basis of its hooked styles, which can be seen clearly here.]]
In 1981, [[Alex George]] published his classic 1981 monograph ''The Genus ''Banksia'' L.f. (Proteaceae)''. George's arrangement was based on a variety of properties including leaf, [[carpel|style]], pollen-presenter, [[follicle (fruit)|follicle]] and seed characters, with the criterion that a taxa was considered a distinct species only if it exhibited a "significant and consistent difference in the morphology of flowers and/or fruit".<ref name="George 1981"/> It was the first thorough revision of the taxonomy of ''Banksia'' for over a century, and formed the basis for George's 1984 ''The Banksia Book'',<ref name="George 1984">{{cite book | first = Alex|last= George | year = 1996 | title = The Banksia Book | publisher = Kangaroo Press and The Society for Growing Australian Plants (NSW Region) | id = ISBN 0-86417-818-2}}</ref> which remains the standard text on the genus, and the treatment of ''Banksia'' in the ''[[Flora of Australia (series)|Flora of Australia]]'' series.<ref name="George 1999"/>
George followed Brown in dividing ''Banksia'' into two subgenera, [[Banksia subg. Banksia|''Banksia'']] and ''Isostylis''. He then divided subgenus ''Banksia'' into two sections: [[Banksia sect. Banksia|''Banksia'']] for species with straight or slightly curved styles, and [[Banksia sect. Oncostylis|''Oncostylis'']] for species with hooked styles.<ref name="George 1981"> </ref> These two sections were then divided into nine and three series respectively. The arrangement into series largely followed Bentham, with series ''Orthostylis'' remaining somewhat heterogeneous, and ''Cyrtostylis'' remaining highly heterogeneous.<ref name="Thiele 1996"/>
The standard taxonomic arrangement of ''Banksia'', as determined by George and published in the ''Flora of Australia'' series, may be summarised as follows:
:'''Genus ''[[Banksia]]'''''
::Subgenus [[Banksia subg. Banksia|''Banksia'']]
:::Section [[Banksia sect. Banksia|''Banksia'']]
::::Series [[Banksia ser. Salicinae|''Salicinae'']]
:::::''[[Banksia dentata|B. dentata]]'' - ''[[Banksia aquilonia|B. aquilonia]]'' - ''[[Banksia integrifolia|B. integrifolia]]'' - ''[[Banksia plagiocarpa|B. plagiocarpa]]'' - ''[[Banksia oblongifolia|B. oblongifolia]]'' - ''[[Banksia robur|B. robur]]'' - ''[[Banksia conferta|B. conferta]]'' - ''[[Banksia paludosa|B. paludosa]]'' - ''[[Banksia marginata|B. marginata]]'' - ''[[Banksia canei|B. canei]]'' - ''[[Banksia saxicola|B. saxicola]]''
::::Series [[Banksia ser. Grandes|''Grandes'']]
:::::''[[Banksia grandis|B. grandis]]'' - ''[[Banksia solandri|B. solandri]]''
::::Series [[Banksia ser. Banksia|''Banksia'']]
:::::''[[Banksia serrata|B. serrata]]'' - ''[[Banksia aemula|B. aemula]]'' - ''[[Banksia ornata|B. ornata]]'' - ''[[Banksia baxteri|B. baxteri]]'' - ''[[Banksia speciosa|B. speciosa]]'' - ''[[Banksia menziesii|B. menziesii]]'' - ''[[Banksia candolleana|B. candolleana]]'' - ''[[Banksia sceptrum|B. sceptrum]]''
::::Series [[Banksia ser. Crocinae|''Crocinae'']]
:::::''[[Banksia prionotes|B. prionotes]]'' - ''[[Banksia burdettii|B. burdettii]]'' - ''[[Banksia hookeriana|B. hookeriana]]'' - ''[[Banksia victoriae|B. victoriae]]''
::::Series [[Banksia ser. Prostratae|''Prostratae'']]
:::::''[[Banksia goodii|B. goodii]]'' - ''[[Banksia gardneri|B. gardneri]]'' - ''[[Banksia chamaephyton|B. chamaephyton]]'' - ''[[Banksia blechnifolia|B. blechnifolia]]'' - ''[[Banksia repens|B. repens]]'' - ''[[Banksia petiolaris|B. petiolaris]]''
::::Series [[Banksia ser. Cyrtostylis|''Cyrtostylis'']]
:::::''[[Banksia media|B. media]]'' - ''[[Banksia praemorsa|B. praemorsa]]'' - ''[[Banksia epica|B. epica]]'' - ''[[Banksia pilostylis|B. pilostylis]]'' - ''[[Banksia attenuata|B. attenuata]]'' - ''[[Banksia ashbyi|B. ashbyi]]'' - ''[[Banksia benthamiana|B. benthamiana]]'' - ''[[Banksia audax|B. audax]]'' - ''[[Banksia lullfitzii|B. lullfitzii]]'' - ''[[Banksia elderiana|B. elderiana]]'' - ''[[Banksia laevigata|B. laevigata]]'' - ''[[Banksia elegans|B. elegans]]'' - ''[[Banksia lindleyana|B. lindleyana]]''
::::Series [[Banksia ser. Tetragonae|''Tetragonae'']]
:::::''[[Banksia lemanniana|B. lemanniana]]'' - ''[[Banksia caleyi|B. caleyi]]'' - ''[[Banksia aculeata|B. aculeata]]''
::::Series ''Bauerinae''
:::::''[[Banksia baueri|B. baueri]]''
::::Series [[Banksia ser. Quercinae|''Quercinae'']]
:::::''[[Banksia quercifolia|B. quercifolia]]'' - ''[[Banksia oreophila|B. oreophila]]''
:::Section ''Coccinea''
:::::''[[Banksia coccinea|B. coccinea]]''
:::Section [[Banksia sect. Oncostylis|''Oncostylis'']]
::::Series [[Banksia ser. Spicigerae|''Spicigerae'']]
:::::''[[Banksia spinulosa|B. spinulosa]]'' - ''[[Banksia ericifolia|B. ericifolia]]'' - ''[[Banksia verticillata|B. verticillata]]'' - ''[[Banksia seminuda|B. seminuda]]'' - ''[[Banksia littoralis|B. littoralis]]'' - ''[[Banksia occidentalis|B. occidentalis]]'' - ''[[Banksia brownii|B. brownii]]''
::::Series ''Tricuspidae''
:::::''[[Banksia tricuspis|B. tricuspis]]''
::::Series ''Dryandroidae''
:::::''[[Banksia dryandroides|B. dryandroides]]''
::::Series [[Banksia ser. Abietinae|''Abietinae'']]
:::::''[[Banksia sphaerocarpa|B. sphaerocarpa]]'' - ''[[Banksia micrantha|B. micrantha]]'' - ''[[Banksia grossa|B. grossa]]'' - ''[[Banksia telmatiaea|B. telmatiaea]]'' - ''[[Banksia leptophylla|B. leptophylla]]'' - ''[[Banksia lanata|B. lanata]]'' - ''[[Banksia scabrella|B. scabrella]]'' - ''[[Banksia violacea|B. violacea]]'' - ''[[Banksia incana|B. incana]]'' - ''[[Banksia laricina|B. laricina]]'' - ''[[Banksia pulchella|B. pulchella]]'' - ''[[Banksia meisneri|B. meisneri]]'' - ''[[Banksia nutans|B. nutans]]''
::Subgenus [[Banksia subg. Isostylis|''Isostylis'']]
:::::''[[Banksia ilicifolia|B. ilicifolia]]'' - ''[[Banksia oligantha|B. oligantha]]'' - ''[[Banksia cuneata|B. cuneata]]''
==Cladistic analyses==
===Thiele and Ladiges===
{{main|Thiele and Ladiges' taxonomic arrangement of Banksia}}
In 1996, [[Kevin Thiele]] and Pauline Ladiges published a cladistic analysis of the ''Banksia'' genus in the journal ''[[Australian Systematic Botany]]''.<ref name="Thiele 1996">{{cite journal | author = [[Kevin Thiele|Thiele, Kevin]] and Pauline Y. Ladiges | year = 1996 | title = A Cladistic Analysis of Banksia (Proteaceae) | journal = Australian Systematic Botany | volume = 9 | issue = 5 | pages = 661–733 | url = http://www.publish.csiro.au/?act=view_file&file_id=SB9960661.pdf | accessdate = 2007-01-10 | doi = 10.1071/SB9960661}}</ref> As their [[cladistics|cladogram]] differed substantially from the current taxonomic arrangement, they published a revised arrangement that accorded better with their results. Four varieties were promoted to species rank: ''B. conferta'' var. ''penicillata'' to ''B. penicillata'' (now [[Banksia conferta subsp. penicillata|''B. conferta'' subsp. ''penicillata'']]); [[Banksia gardneri var. brevidentata|''B. gardneri'' var. ''brevidentata'']] to ''B. brevidentata''; [[Banksia gardneri var. hiemalis|''B. gardneri'' var. ''hiemalis'']] to ''B. hiemalis''; and [[Banksia sphaerocarpa var. dolichostyla|''B. sphaerocarpa'' var. ''dolichostyla'']] to ''B. dolichostyla''. Two new series and elevensubseries were introduced; [[Banksia sect. Oncostylis|''Banksia'' sect. ''Oncostylis'']] and [[Banksia ser. Crocinae|''Banksia'' ser. ''Crocinae'']] were discarded; and [[Banksia ser. Cyrtostylis|''Banksia'' ser. ''Cyrtostylis'']] was largely redefined. Six species were left ''[[incertae sedis]]''.<ref name="Thiele 1996"/>
Most aspects of Thiele and Ladiges' arrangement were not accepted by George in his 1999 revision. He stated that "the infrageneric classification and systematic sequence presented here are modified from that of George (1981) and take into account new data revealed in the work of Thiele & Ladiges (1996)", but none of the four promotions to species rank was accepted, and none of the thirteen infrageneric taxa introduced by Thiele and Ladiges was retained.<ref name="George 1999">{{cite encyclopedia | last = George | first = Alex S. | authorlink = Alex George | year = 1999 | title = Banksia | editor = Wilson, Annette (ed.) | encyclopedia = Flora of Australia | volume = Volume 17B: Proteaceae 3: Hakea to Dryandra | pages = 175–251 | location = Collingwood, Victoria | publisher = CSIRO Publishing / Australian Biological Resources Study | id = ISBN 0-643-06454-0}}</ref> However a number of Australian [[herbarium|herbaria]] have continued to follow Thiele and Ladiges on some points, for example by recognising the four species that they promoted.<ref name="Olde 2002">{{cite journal | first= Peter| last=Olde|coauthors=Neil R. Marriott| year = 2002 | title = One new ''Banksia'' and two new ''Grevillea'' species (Proteaceae: Grevilleoideae) from Western Australia | journal = Nuytsia | volume = 15 | issue = 1 | pages = 85–99 | url = http://users.bigpond.net.au/macarthuraps/new%20grevilleas.pdf | accessdate = 2007-01-10}}</ref>
===DNA Analysis===
Following on from an earlier molecular study,<ref>Mast AR. (1998) Molecular systematics of subtribe Banksiinae (Banksia and Dryandra; Proteaceae) based on cpDNA and nrDNA sequence data: implications for taxonomy and biogeography. ''Australian Systematic Botany'' '''11''': 321-342</ref> [[Austin Mast]] and co-authors published cladistic analyses of genetic data from [[DNA]] samples of almost all species of ''Banksia'', along with five ''Dryandra'' species in 2002 and 2005. Their results indicated the presence of two large clades of ''Banksia'', which they named "'''/Cryptostomata'''" ("hidden stomates") and "'''/Phanerostomata'''" ("visible stomates").
The '''/Phanerostomata''' were defined as those taxa in which the leaf [[stoma]]ta occur superficially or in shallow pits. These taxa are typically tall shrubs and trees that occur in moist areas; they have unbeaked follicles and soft, short-lived leaves that are in many cases needle-like. The clade includes all eastern taxa of the series Salicinae and Spicigerae (that is, all taxa except ''B. serrata'', ''B. aemula'' and ''B. ornata'') and also the western Spicigerae, Quercinae, Grandes, Abietinae and Dryandroideae.
The '''/Cryptostomata''' were defined as those taxa in which the leaf stomata occur in crypts with constricted entrances. These are usually small shrubs that occur on dry, infertile sandplains. They have beaked follicles and thick, tough, long-lived serrated leaves. It includes all other western taxa, plus the eastern species ''B. serrata'', ''B. aemula'' and ''B. ornata'', and also appears to include ''Dryandra''. It is worth noting that ''Dryandra'' does not appear especially closely related to the ''Isostylis'' group, which is instead most closely related to ''B. elegans'' and then ''[[Banksia attenuata|B. attenuata]]''.
Thus, the results presented by Mast ''et al.'' strongly suggest that ''Banksia'' is polyphyletic with ''Dryandra''. Although they did not propose a new taxonomic arrangement, they did consider various options for adjusting the current accepted arrangement to remove the polyphyly. They conclude that the simplest and least disruptive solution would be to merge ''Dryandra'' into ''Banksia''.<ref name="Mast 2002"/><ref name="Mast 2005"/>
==References==
{{wikisourcelang|la|Supplementum Plantarum/Index Generum/Banksia|Supplementum Plantarum, Index Generum, 1362. Banksia}}
{{wikisourcepar|Flora Australiensis/Volume V/CIV. Proteaceae/28. Banksia}}
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<references/>
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[[Category:Taxonomy of Banksia| ]]
[[fr:Taxonomie du genre Banksia]]