Banksia subg. Banksia

taxobox
name = "Banksia" subg. "Banksia"

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image_caption = "Banksia serrata" (Saw Banksia), the type species of "Banksia".
regnum = Plantae
unranked_divisio = Angiosperms
unranked_classis = Eudicots
ordo = Proteales
familia = Proteaceae
genus = "Banksia"
subgenus = "Banksia" L.f. subg. "Banksia"|

"Banksia" subg. "Banksia" is a valid botanic name for a subgenus of "Banksia". As an autonym, it necessarily contains the type species 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". Under Brown's arrangement, "Banksia" was divided into two groups based on inflorescence 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".cite book | author = Brown, Robert | year = 1810 | title = Prodromus Florae Novae Hollandiae et Insulae Van Diemen | publisher = Taylor | location = London]

Brown published a further eleven species in 1830, placing all of them in "Banksia verae". The 1830 circumscription of "Banksia verae" was as follows:cite book | author = Brown, Robert | year = 1830 | title = Supplementum Primum Prodromi Florae Novae Hollandiae | publisher = Taylor | location = London] :"Banksia"::"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" by Stephan Endlicher in his 1847 fourth supplement to "Genera Plantarum Secundum Ordines Naturales Disposita",cite book | first = Stephan | last = Endlicher | authorlink = Stephan Endlicher | year = 1847 | title = Genera Plantarum Secundum Ordines Naturales Disposita Supplement 4 | pages = [http://www.botanicus.org/page/503709 88] ] and this name was retained in Carl Meissner'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".cite encyclopedia | author = Meissner, Carl | 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] 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::"Banksia"::"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 in 1870. Instead, Bentham divided the genus into five section, retaining "B." sect. "Isostylis", but dividing the remaining species into four sections.cite encyclopedia | author = Bentham, George | year = 1870 | title = Banksia | encyclopedia = Flora Australiensis | volume = Volume 5: Myoporineae to Proteaceae | pages = 541–562 | location = London | publisher = L. Reeve & Co.]

"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)". George reinstated Meissner's "Eubanksia" and "Isostylis", giving them subgeneric rank. By this time, however, the rules of botanical nomenclature had been formalised in such a way that "Eubanksia" was required to take the autonym "B." subg. "Banksia".The genus Banksia L.f. (Proteaceae)]

George's arrangement was overturned in 1996 by Kevin Thiele and Pauline Ladiges, but in 1999 George published a slightly modified version in his treatment of "Banksia" for the "Flora of Australia" series of monographs. George's 1999 arrangement is not universally accepted, and accords poorly with recently published cladistic analyses, yet it remains the most recently published arrangement.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]

In George's taxonomic arrangement of "Banksia", "B." subg. "Banksia" is further divided into three sections, primarily on the shape of the style. "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". "B." sect. "Oncostylis" contains those species whose styles are hooked; it contains about 20 species, which are further divided into four series. Its is circumscribed as follows::"Banksia"::"B." subg. "Banksia":::"B." sect. "Banksia"::::"B." ser. "Salicinae":::::"B. dentata":::::"B. aquilonia":::::"B. integrifolia"::::::"B. integrifolia" subsp. "integrifolia"::::::"B. integrifolia" subsp. "compar"::::::"B. integrifolia" subsp. "monticola":::::"B. plagiocarpa":::::"B. oblongifolia":::::"B. robur":::::"B. conferta"::::::"B. conferta" subsp. "conferta"::::::"B. conferta" subsp. "penicillata":::::"B. paludosa"::::::"B. paludosa" subsp. "astrolux"::::::"B. paludosa" subsp. "paludosa":::::"B. marginata":::::"B. canei":::::"B. saxicola::::"B." ser. "Grandes":::::"B. grandis":::::"B. solandri"::::"B." ser. "Banksia":::::"B. serrata":::::"B. aemula":::::"B. ornata":::::"B. baxteri":::::"B. speciosa":::::"B. menziesii":::::"B. candolleana":::::"B. sceptrum"::::"B." ser. "Crocinae":::::"B. prionotes":::::"B. burdettii":::::"B. hookeriana":::::"B. victoriae"::::"B." ser. "Prostratae":::::"B. goodii":::::"B. gardneri"::::::"B. gardneri" var. "gardneri"::::::"B. gardneri" var. "brevidentata"::::::"B. gardneri" var. "hiemalis":::::"B. chamaephyton":::::"B. blechnifolia":::::"B. repens":::::"B. petiolaris"::::"B." ser. "Cyrtostylis":::::"B. media":::::"B. praemorsa":::::"B. epica":::::"B. pilostylis":::::"B. attenuata":::::"B. ashbyi":::::"B. benthamiana":::::"B. audax":::::"B. lullfitzii":::::"B. elderiana":::::"B. laevigata"::::::"B. laevigata" subsp. "laevigata"::::::"B. laevigata" subsp. "fuscolutea":::::"B. elegans":::::"B. lindleyana"::::"B." ser. "Tetragonae":::::"B. lemanniana":::::"B. caleyi":::::"B. aculeata"::::"B." ser. "Bauerinae":::::"B. baueri"::::"B." ser. "Quercinae":::::"B. quercifolia":::::"B. oreophila":::"B." sect. "Coccinea":::::"B. coccinea":::"B." sect. "Oncostylis"::::"B." ser. "Spicigerae":::::"B. spinulosa"::::::"B. spinulosa" var. "spinulosa"::::::"B. spinulosa" var. "collina"::::::"B. spinulosa" var. "neoanglica"::::::"B. spinulosa" var. "cunninghamii":::::"B. ericifolia"::::::"B. ericifolia" subsp. "ericifolia"::::::"B. ericifolia" subsp. "macrantha":::::"B. verticillata":::::"B. seminuda":::::"B. littoralis":::::"B. occidentalis":::::"B. brownii"::::"B." ser. "Tricuspidae":::::"B. tricuspis"::::"B." ser. "Dryandroideae":::::"B. dryandroides"::::"B." ser. "Abietinae":::::"B. sphaerocarpa"::::::"B. sphaerocarpa" var. "sphaerocarpa"::::::"B. sphaerocarpa" var. "caesia"::::::"B. sphaerocarpa" var. "dolichostyla":::::"B. micrantha":::::"B. grossa":::::"B. telmatiaea":::::"B. leptophylla"::::::"B. leptophylla" var. "leptophylla"::::::"B. leptophylla" var. "melletica":::::"B. lanata":::::"B. scabrella":::::"B. violacea":::::"B. incana":::::"B. laricina":::::"B. pulchella":::::"B. meisneri"::::::"B. meisneri" subsp. "meisneri"::::::"B. meisneri" subsp. "ascendens":::::"B. nutans"::::::"B. nutans" var. "nutans"::::::"B. nutans" var. "cernuella"::"B." subg. "Isostylis" (3 species)

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

In 1996, Kevin Thiele and Pauline Ladiges published the results of a cladistic analysis of "Banksia". They found George's arrangement to accord fairly closely with their inferred cladogram, 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 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 unless "B. elegans" (Elegant Banksia) was excluded. They ended up listing "B. elegans" and five other species as "incertae sedis", but otherwise maintained George's circumscription of the subgenus.cite journal |first=Kevin|last=Thiele| coauthors = Pauline Y. 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]

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::"Banksia"::"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." subg. "Banksia" "sensu" Mast and Thiele

In 2005, Austin Mast, Eric Jones and Shawn Havery published the results of their cladistic analyses of DNA sequence data for "Banksia". They inferred a phylogeny very greatly different from the accepted taxonomic arrangement, including finding "Banksia" to be paraphyletic with respect to "Dryandra".cite journal | author = Mast, Austin R. | 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 | pages = 321–342 | doi = 10.1071/SB97026] cite journal | author = Mast, Austin R. and 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 = 1311–1323 | 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] cite journal | author = 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 | doi = 10.1071/SB04015] 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 cotyledons. 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.cite journal | author = Mast, Austin R. and Kevin Thiele | 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]

References

External links

*Flora of Australia Online|name="Banksia" subg. "Banksia"|id=3382