''Banksia'' subg. ''Banksia'' is a valid

botanic name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

for a

subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...

of ''

Banksia ''Banksia'' is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes, and fruiting "cones" and heads. ''Banksias'' range i ...

''. As an

autonym Autonym may refer to: * Autonym, the name used by a person to refer to themselves or their language; see Exonym and endonym * Autonym (botany), an automatically created infrageneric or infraspecific name See also * Nominotypical subspecies, in zo ...

, it necessarily contains the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

of ''Banksia'', '' B. serrata'' (Saw Banksia). Within this constraint, however, there have been various circumscriptions.

''Banksia verae''

''B.'' subg. ''Banksia'' can be traced back to ''Banksia verae'', an unranked taxon published by Robert Brown in his 1810 ''

Prodromus Florae Novae Hollandiae et Insulae Van Diemen ''Prodromus Florae Novae Hollandiae et Insulae Van Diemen'' (Prodromus of the Flora of New Holland and Van Diemen's Land) is a flora of Australia written by botanist Robert Brown and published in 1810. Often referred to as ''Prodromus Flora Novae ...

''. Under Brown's arrangement, ''Banksia'' was divided into two groups based on

inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

shape. ''Banksia verae'' was defined as containing those ''Banksia'' taxa with the elongate flower spike typical of ''Banksia'', and it thus contained all but one species. The remaining species, '' B. ilicifolia'' (Holly-leaved Banksia), has a dome-shaped head and so was placed alone in '' Isostylis''. Brown published a further eleven species in 1830, placing all of them in ''Banksia verae''. The 1830 circumscription of ''Banksia verae'' was as follows: :''

'' ::''Banksia verae'' :::'' B. pulchella'' :::'' B. sphærocarpa'' :::'' B. nutans'' :::'' B. ericifolia'' :::'' B. spinulosa'' :::''B. Cunninghamii'' (now ''B. spinulosa'' var. ''cunninghamii'') :::''B. collina'' (now ''B. spinulosa'' var. ''collina'') :::'' B. occidentalis'' :::'' B. littoralis'' :::'' B. marginata'' :::''B. depressa'' (now '' B. marginata'') :::''B. patula'' (now '' B. marginata'') :::''B. australis'' (now '' B. marginata'') :::''B. insularis'' (now '' B. marginata'') :::'' B. integrifolia'' :::''B. compar'' (now ''B. integrifolia'' subsp. ''compar'') :::'' B. verticillata'' :::'' B. coccinea'' :::'' B. paludosa'' :::'' B. oblongifolia'' :::''B. latifolia'' (now '' B. robur'') :::''B. marcescens'' (now '' B. praemorsa'') :::'' B. media'' :::'' B. attenuata'' :::'' B. Caleyi'' :::'' B. Baueri'' :::'' B. Menziesii'' :::''B. elatior'' (now '' B. aemula'') :::'' B. serrata'' :::'' B. æmula'' :::'' B. dentata'' :::'' B. quercifolia'' :::'' B. speciosa'' :::'' B. Solandri'' :::'' B. grandis'' :::'' B. Baxteri'' :::'' B. Goodii'' :::''B. prostrata'' (now '' B. gardneri'') :::'' B. repens'' :::'' B. Dryandroides'' :::'' B. Brownii'' ::'' Isostylis'' (one species)

''Eubanksia''

''Banksia verae'' was renamed ''

Eubanksia ''Banksia'' sect. ''Eubanksia'' is an obsolete section of ''Banksia''. There have been two circumscriptions, one of which is synonymous with the recently abandoned ''B.'' subg. ''Banksia'' ''sensu'' Alex George, the other having no modern equi ...

'' by

Stephan Endlicher Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian botanist, numismatist and Sinologist. He was a director of the Botanical Garden of Vienna. Bio ...

in his 1847 fourth supplement to '' Genera Plantarum Secundum Ordines Naturales Disposita'', and this name was retained in

Carl Meissner Carl Daniel Friedrich Meissner (1 November 1800 – 2 May 1874) was a Swiss botanist. Biography Born in Bern, Switzerland on 1 November 1800, he was christened Meisner but later changed the spelling of his name to Meissner. For most of his 40 ...

's 1856 arrangement. Meissner gave ''Eubanksia'' sectional rank, further dividing it into four series, ''B.'' ser. ''Abietinæ'', ''B.'' ser. ''Salicinae'', ''B.'' ser. ''Quercinae'' and ''B.'' ser. ''Dryandroideae''. These series were defined in terms of leaf shape alone, and were hence all highly heterogeneous, with the exception of ''B.'' ser. ''Abietinæ'', which contained only (but not all) species with hooked styles. Meissner's 1856 circumscription of ''B.'' sect. ''Eubanksia'' was as follows: :''

'' :: ''B.'' sect. ''Eubanksia'' ::: ''B.'' ser. ''Abietinæ'' ::::'' B. pulchella'' ::::'' B. Meisneri'' ::::'' B. sphærocarpa'' :::::''B. sphaerocarpa'' var. ''glabrescens'' (now '' B. incana'') ::::''B. pinifolia'' (now '' B. leptophylla'') ::::'' B. nutans'' ::::'' B. ericifolia'' ::::'' B. spinulosa'' ::::'' B. tricuspis'' ::: ''B.'' ser ''Salicinæ'' ::::''B. cunninghamii'' (now ''B. spinulosa'' var. ''cunninghamii'') ::::''B. collina'' (now ''B. spinulosa'' var. ''collina'') ::::'' B. occidentalis'' ::::'' B. littoralis'' ::::''B. cylindrostachya'' (now '' B. attenuata'') ::::'' B. lindleyana'' ::::'' B. marginata'' :::::''B. marginata'' var. ''Cavanillesii'' (now '' B. marginata'') :::::''B. marginata'' var. ''microstachya'' (now '' B. marginata'') :::::''B. marginata'' var. ''humilis'' (now '' B. marginata'') ::::''B. depressa'' (now '' B. marginata'') :::::''B. depressa'' var. ''subintegra'' (now '' B. marginata'') ::::''B. patula'' (now '' B. marginata'') ::::''B. australis'' (now '' B. marginata'') ::::''B. Gunnii'' (now '' B. marginata'') ::::''B. insularis'' (now '' B. marginata'') ::::'' B. integrifolia'' :::::''B. integrifolia'' var. ''minor'' (now ''B. integrifolia'' subsp. ''integrifolia'') :::::''B. integrifolia'' var. ''major'' (now ''B. integrifolia'' subsp. ''integrifolia'') :::::''B. integrifolia'' var. ''dentata'' (now ''B. robur'') ::::''B. compar'' (now ''B. integrifolia'' subsp. ''compar'') ::::'' B. paludosa'' ::::'' B. verticillata'' ::::'' B. media'' ::::'' B. attenuata'' ::::''B. elatior'' (now '' B. aemula'') ::::'' B. lævigata'' ::::'' B. Hookeriana'' ::::'' B. prionotes'' ::::'' B. Menziesii'' ::: ''B.'' ser. ''Quercinæ'' ::::'' B. coccinea'' ::::'' B. sceptrum'' ::::'' B. Baueri'' ::::'' B. ornata'' ::::''B. latifolia'' (now '' B. robur'') ::::''B. marcescens'' (now '' B. praemorsa'') ::::'' B. oblongifolia'' ::::'' B. serrata'' ::::'' B. æmula'' ::::'' B. Caleyi'' :::::''B. caleyi'' var. ''sinuosa'' (now '' B. caleyi'') ::::'' B. Lemanniana'' ::::'' B. quercifolia'' ::::'' B. dentata'' ::::''B. prostrata'' (now '' B. gardneri'') ::::'' B. Goodii'' ::::''B. barbigera'' ::::'' B. repens'' ::::'' B. Solandri'' :::::''B. solandri'' var. ''major'' (now '' B. solandri'') ::: ''B.'' ser. ''Dryandroideæ'' ::::'' B. grandis'' ::::'' B. Baxteri'' ::::'' B. speciosa'' ::::'' B. Victoriæ'' ::::'' B. elegans'' ::::'' B. Candolleana'' ::::'' B. dryandroides'' ::::'' B. Brownii'' :: ''B.'' sect. ''Isostylis'' (one species) The top-level split into ''Eubanksia'' and ''Isostylis'' was abandoned by

George Bentham George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studi ...

in 1870. Instead, Bentham divided the genus into five section, retaining ''B.'' sect. ''Isostylis'', but dividing the remaining species into four sections.

''B.'' subg. ''Banksia'' ''sensu'' George

In 1981, Alex George published a thorough revision of ''Banksia'' in his classic monograph ''

The genus Banksia L.f. (Proteaceae) "The genus ''Banksia'' L.f. (Proteaceae)" is a 1981 monograph by Alex George on the taxonomy of the plant genus ''Banksia''. Published by the Western Australian Herbarium as ''Nuytsia'' 3(3), it presented George's taxonomic arrangement of ''B ...

''. George reinstated Meissner's ''Eubanksia'' and ''Isostylis'', giving them subgeneric rank. By this time, however, the rules of

botanical nomenclature Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from Alpha taxonomy, taxonomy. Plant taxonomy is concerned with grouping and classifying plants; botanical nomenclature then provides names for the ...

had been formalised in such a way that ''Eubanksia'' was required to take the

''B.'' subg. ''Banksia''. George's arrangement was overturned in 1996 by

Kevin Thiele Kevin R. Thiele is currently an adjunct associate professor at the University of Western Australia and the director of Taxonomy Australia. He was the curator of the Western Australian Herbarium from 2006 to 2015. His research interests include ...

and

Pauline Ladiges Pauline Yvonne Ladiges (born 1948) is a botanist whose contributions have been significant both in building the field of taxonomy, ecology and historical biogeography of Australian plants, particularly Eucalypts and flora, and in science educa ...

, but in 1999 George published a slightly modified version in his treatment of ''Banksia'' for the ''

Flora of Australia The flora of Australia comprises a vast assemblage of plant species estimated to over 30,000 vascular and 14,000 non-vascular plants, 250,000 species of fungi and over 3,000 lichens. The flora has strong affinities with the flora of Gondwana, ...

'' series of monographs. George's 1999 arrangement is not universally accepted, and accords poorly with recently published

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

analyses, yet it remains the most recently published arrangement. In George's taxonomic arrangement of ''Banksia'', ''B.'' subg. ''Banksia'' is further divided into three sections, primarily on the shape of the

style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...

. ''B.'' sect. ''Banksia'' species have a straight or curved, but never hooked, style; this section contains about 50 species that are further divided into nine series. ''B.'' sect. ''Coccinea'' contains a single species, ''

Banksia coccinea ''Banksia coccinea'', commonly known as the scarlet banksia, waratah banksia or Albany banksia, is an erect shrub or small tree in the family Proteaceae. Its distribution in the wild is along the south west coast of Western Australia, from Denm ...

''. ''B.'' sect. ''Oncostylis'' contains those species whose styles are hooked; it contains about 20 species, which are further divided into four series. It is circumscribed as follows: :''

''B.'' subg. ''Banksia'' ''sensu'' Thiele and Ladiges

In 1996,

and Pauline Ladiges published the results of a

analysis of ''Banksia''. They found George's arrangement to accord fairly closely with their inferred

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

, so sought to publish a taxonomic arrangement that reflected their phylogeny whilst being minimally disruptive to the then-current arrangement. They accepted both of George's subgenera prior to the analysis, using each as an

outgroup Outgroup may refer to: * Outgroup (cladistics), an evolutionary-history concept * Outgroup (sociology), a social group {{disambig ...

in the analysis of the other. Thus their analysis yielded little information about the circumscription and placement of ''B.'' subg. ''Banksia''. They did find, however, that the subgenus was not

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

unless '' B. elegans'' (Elegant Banksia) was excluded. They ended up listing ''B. elegans'' and five other species as ''

incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...

'', but otherwise maintained George's circumscription of the subgenus. In Thiele and Ladiges' taxonomic arrangement of ''Banksia'', ''B.'' subg. ''Banksia'' is divided into twelve series. Its placement and circumscription may be summarised as follows: :''

'' :: ''B.'' subg. ''Isostylis'' (three species) ::'' B. elegans'' (''incertae sedis'') ::''B.'' subg. ''Banksia'' ::: ''B.'' ser. ''Tetragonae'' ::::'' B. elderiana'' ::::'' B. lemanniana'' ::::'' B. caleyi'' ::::'' B. aculeata'' ::: ''B.'' ser. ''Lindleyanae'' ::::'' B. lindleyana'' ::: ''B.'' ser. ''Banksia'' :::: ''B.'' subser. ''Banksia'' :::::'' B. ornata'' :::::'' B. serrata'' :::::'' B. aemula'' :::: ''B.'' subser. ''Cratistylis'' :::::'' B. candolleana'' :::::'' B. sceptrum'' :::::'' B. baxteri'' :::::'' B. speciosa'' :::::'' B. menziesii'' :::::'' B. burdettii'' :::::'' B. victoriae'' :::::'' B. hookeriana'' :::::'' B. prionotes'' :::'' B. baueri'' (''incertae sedis'') :::'' B. lullfitzii'' (''incertae sedis'') :::'' B. attenuata'' (''incertae sedis'') :::'' B. ashbyi'' (''incertae sedis'') :::'' B. coccinea'' (''incertae sedis'') ::: ''B.'' ser. ''Prostratae'' ::::'' B. petiolaris'' ::::'' B. repens'' ::::'' B. chamaephyton'' ::::'' B. blechnifolia'' ::::''B. hiemalis'' (now ''B. gardneri'' var. ''hiemalis'') ::::'' B. gardneri'' ::::''B. brevidentata'' (now ''B. gardneri'' var. ''brevidentata'') ::::'' B. goodii'' ::: ''B.'' ser. ''Cyrtostylis'' ::::'' B. pilostylis'' ::::'' B. media'' ::::'' B. epica'' ::::'' B. praemorsa'' ::: ''B.'' ser. ''Ochraceae'' ::::'' B. benthamiana'' ::::'' B. audax'' ::::'' B. laevigata'' ::::: ''B. laevigata'' subsp. ''laevigata'' ::::: ''B. laevigata'' subsp. ''fuscolutea'' ::: ''B.'' ser. ''Grandes'' ::::'' B. grandis'' ::::'' B. solandri'' ::: ''B.'' ser. ''Salicinae'' :::: ''B.'' subser. ''Acclives'' :::::'' B. oblongifolia'' :::::'' B. plagiocarpa'' :::::'' B. robur'' :::::'' B. dentata'' :::: ''B.'' subser. ''Integrifoliae'' :::::'' B. marginata'' :::::'' B. conferta'' :::::''B. penicillata'' (now ''B. conferta'' subsp. ''penicillata'') :::::'' B. paludosa'' :::::'' B. canei'' :::::'' B. saxicola'' :::::'' B. integrifolia'' :::::: ''B. integrifolia'' subsp. ''integrifolia'' :::::: ''B. integrifolia'' subsp. ''monticola'' :::::: ''B. integrifolia'' subsp. ''compar'' ::::::''B. integrifolia'' subsp. ''aquilonia'' (now '' B. aquilonia'') ::: ''B.'' ser. ''Spicigerae'' :::: ''Banksia'' subser. ''Spinulosae'' :::::'' B. spinulosa'' :::::: ''B. spinulosa'' var. ''spinulosa'' :::::: ''B. spinulosa'' var. ''collina'' :::::: ''B. spinulosa'' var. ''cunninghamii'' :::::: ''B. spinulosa'' var. ''neoanglica'' :::: ''B.'' subser. ''Ericifoliae'' :::::'' B. ericifolia'' :::::: ''B. ericifolia'' var. ''ericifolia'' :::::: ''B. ericifolia'' var. ''macrantha'' :::: ''B.'' subser. ''Occidentales'' :::::'' B. occidentalis'' :::::'' B. brownii'' :::::'' B. seminuda'' :::::'' B. verticillata'' :::::'' B. littoralis'' ::: ''B.'' ser. ''Quercinae'' ::::'' B. quercifolia'' ::::'' B. oreophila'' ::: ''B.'' ser. ''Dryandroideae'' ::::'' B. dryandroides'' ::: ''B.'' ser. ''Abietinae'' :::: ''B.'' subser. ''Nutantes'' :::::'' B. nutans'' :::::: ''B. nutans'' var. ''nutans'' :::::: ''B. nutans'' var. ''cernuella'' :::: ''B.'' subser. ''Sphaerocarpae'' :::::'' B. grossa'' :::::''B. dolichostyla'' (now ''B. sphaerocarpa'' var. ''dolichostyla'') :::::'' B. micrantha'' :::::'' B. sphaerocarpa'' :::::: ''B. sphaerocarpa'' var. ''sphaerocarpa'' :::::: ''B. sphaerocarpa'' var. ''caesia'' :::: ''B.'' subser. ''Leptophyllae'' :::::'' B. telmatiaea'' :::::'' B. scabrella'' :::::'' B. leptophylla'' :::::: ''B. leptophylla'' var. ''melletica'' :::::: ''B. leptophylla'' var. ''leptophylla'' :::::'' B. lanata'' :::: ''B.'' subser. ''Longistyles'' :::::'' B. violacea'' :::::'' B. laricina'' :::::'' B. incana'' :::::'' B. tricuspis'' :::::'' B. pulchella'' :::::'' B. meisneri'' :::::: ''B. meisneri'' var. ''meisneri'' :::::: ''B. meisneri'' var. ''ascendens'': :: ''B.'' ser. ''Dryandra''

''B.'' subg. ''Banksia'' ''sensu'' Mast and Thiele

In 2005,

Austin Mast Austin R. Mast is a research botanist. Born in 1972, he obtained a Ph.D. from the University of Wisconsin–Madison in 2000. He is currently a professor within the Department of Biological Science at Florida State University (FSU), and has been dire ...

, Eric Jones and Shawn Havery published the results of their cladistic analyses of DNA sequence data for ''Banksia''. They inferred a

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

very greatly different from the accepted taxonomic arrangement, including finding ''Banksia'' to be

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

with respect to ''

Dryandra ''Banksia'' ser. ''Dryandra'' is a series of 94 species of shrub to small tree in the plant genus ''Banksia''. It was considered a separate genus named ''Dryandra'' until early 2007, when it was merged into ''Banksia'' on the basis of extensiv ...

''. 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 ''B.'' subg. ''Spathulatae'' for the species having spoon-shaped

cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...

s. Thus ''B.'' subg. ''Banksia'' was redefined as containing the species lacking spoon-shaped cotyledons. Mast and Thiele have foreshadowed publishing a full arrangement once DNA sampling of ''Dryandra'' is complete.

References

External links

* {{DEFAULTSORT:Banksia subg. Banksia subg. Banksia Plant subgenera