Bentham's taxonomic arrangement of Dryandra
Encyclopedia
George Bentham
George Bentham
George Bentham CMG FRS was an English botanist, characterized by Duane Isely as "the premier systematic botanist of the nineteenth century".- Formative years :...

's taxonomic arrangement of Dryandra (now B. ser. Dryandra) was published in 1870, in Volume 5 of Bentham's Flora Australiensis
Flora Australiensis
Flora Australiensis: a description of the plants of the Australian Territory, more commonly referred to as Flora Australiensis, and also known by its standard abbreviation Fl. Austral., is a seven-volume flora of Australia published between 1863 and 1878 by George Bentham, with the assistance of...

. It replaced the 1856 arrangement
Meissner's taxonomic arrangement of Dryandra
Carl Meissner's taxonomic arrangement of Dryandra, now Banksia ser. Dryandra, was published in 1856 as part of his chapter on the Proteaceae in A. P. de Candolle's Prodromus systematis naturalis regni vegetabilis...

 of Carl Meissner
Carl Meissner
Carl Daniel Friedrich Meissner was a Swiss botanist.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 year career he was Professor of Botany at University of Basel...

, and stood for over a century before being replaced by the 1996 arrangement
George's taxonomic arrangement of Dryandra
Alex George's taxonomic arrangement of Dryandra was the first modern-day arrangement of that taxon. First published in Nuytsia in 1996, it superseded the arrangement of George Bentham, which had stood for over a hundred years; it would later form the basis for George's 1999 treatment of Dryandra...

 of Alex George
Alex George
Alexander Segger George is a Western Australian botanist. He is the authority on the plant genera Banksia and Dryandra...

.

Background

The dryandras are a group of proteaceous
Proteaceae
Proteaceae is a family of flowering plants distributed in the Southern Hemisphere. The family comprises about 80 genera with about 1600 species. Together with the Platanaceae and Nelumbonaceae they make up the order Proteales. Well known genera include Protea, Banksia, Embothrium, Grevillea,...

 shrubs endemic to southwest Western Australia
Western Australia
Western Australia is a state of Australia, occupying the entire western third of the Australian continent. It is bounded by the Indian Ocean to the north and west, the Great Australian Bight and Indian Ocean to the south, the Northern Territory to the north-east and South Australia to the south-east...

. For nearly two hundred years they were considered a separate genus, having been published at that rank in 1810 by Robert Brown
Robert Brown (botanist)
Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...

. In 2007 it was transferred into the genus Banksia as B. ser. Dryandra. There are now just under 100 species, plus numerous subspecies and varieties.

The first infrageneric arrangement of Dryandra was Brown's 1810 arrangement
Brown's taxonomic arrangement of Dryandra
Robert Brown's taxonomic arrangement of Dryandra was the first arrangement of what is now Banksia ser. Dryandra. His initial arrangement was published in 1810, and a further arrangement, including an infrageneric classification, followed in 1830...

, which listed 13 species, but did not attempt an infrageneric classification. Twenty years later, Brown published a revised arrangement
Brown's taxonomic arrangement of Dryandra
Robert Brown's taxonomic arrangement of Dryandra was the first arrangement of what is now Banksia ser. Dryandra. His initial arrangement was published in 1810, and a further arrangement, including an infrageneric classification, followed in 1830...

 which divided 23 recognised species in three subgenera, and placed one further species in a separate genus Hemiclidia. This was followed by the 1856 arrangement of
Meissner's taxonomic arrangement of Dryandra
Carl Meissner's taxonomic arrangement of Dryandra, now Banksia ser. Dryandra, was published in 1856 as part of his chapter on the Proteaceae in A. P. de Candolle's Prodromus systematis naturalis regni vegetabilis...

 Carl Meissner
Carl Meissner
Carl Daniel Friedrich Meissner was a Swiss botanist.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 year career he was Professor of Botany at University of Basel...

, which recognised 53 species and six varieties, dividing them into three sections and eight series, and maintaining the monospecific genus Hemiclidia.

Bentham's arrangement

Bentham's arrangement was published in 1870, in Volume V of Flora Australiensis
Flora Australiensis
Flora Australiensis: a description of the plants of the Australian Territory, more commonly referred to as Flora Australiensis, and also known by its standard abbreviation Fl. Austral., is a seven-volume flora of Australia published between 1863 and 1878 by George Bentham, with the assistance of...

. He reduced the number of species to 47, and the number of varieties to two, proposing an arrangement consisting of two sections and seven series. Hemiclidia was transferred back into Dryandra.

George Bentham's 1870 taxonomic arrangement of Dryandra may be summarised as follows:
Genus Dryandra (now Banksia ser. Dryandra)
Section Eudryandra
Series Armatæ
Dryandra ser. Armatae
Dryandra ser. Armatae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into Banksia.-According...

D. quercifolia (now B. heliantha
Banksia heliantha
Banksia heliantha, commonly known as Oak-leaved Dryandra, is a shrub endemic to Western Australia.It was known as Dryandra quercifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. praemorsa (now B. undata
Banksia undata
Banksia undata, commonly known as Urchin Dryandra, is a shrub endemic to Western Australia.It was known as Dryandra praemorsa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. cuneata (now B. obovata
Banksia obovata
Banksia obovata, commonly known as Wedge-leaved Dryandra, is a shrub endemic to Western Australia.It was known as Dryandra cuneata until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. falcata (now B. falcata
Banksia falcata
Banksia falcata, commonly known as Prickly Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra falcata until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. armata (now B. armata
Banksia armata
Banksia armata, commonly known as Prickly Dryandra, is a shrub endemic to Western Australia.-Description:B. armata grows as a spreading or upright shrub, up to three metres in height. It has deeply serrated leaves. Its inflorescences are usually bright yellow, but may be pink.-Distribution and...

)
D. longifolia (now B. prolata
Banksia prolata
Banksia prolata is a shrub endemic to Western Australia.It was known as Dryandra longifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. Fraseri (now B. fraseri
Banksia fraseri
Banksia fraseri is a shrub endemic to Western Australia. It was known as Dryandra fraseri until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-Description:...

)
Series Floribundæ
Dryandra ser. Floribundae
Dryandra ser. Floribundae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into...

D. floribunda (now B. sessilis var. sessilis
Banksia sessilis var. sessilis
Banksia sessilis var. sessilis is the most populous and most widespread variety of Banksia sessilis .-Description:The leaves of this variety are larger than those of B. sessilis var. cygnorum but smaller than those of B. sessilis var. cordata. It differs from B. sessilis var...

)
D. floribunda var. major (now B. sessilis var. cordata
Banksia sessilis var. cordata
Banksia sessilis var. cordata is a variety of Banksia sessilis , with unusually large leaves and flower heads. It is a rare variety that is restricted to the extreme south-west corner of Western Australia.-Description:...

)
D. carduacea (now B. squarrosa subsp. squarrosa
Banksia squarrosa subsp. squarrosa
Banksia squarrosa subsp. squarrosa is a subspecies of Banksia squarrosa. As an autonym, it is defined as encompassing the type material of the species. It was known as Dryandra squarrosa subsp. squarrosa until 2007, when Austin Mast and Kevin Thiele sunk all Dryandra into Banksia. As with other...

)
D. carlinoides (now B. carlinoides
Banksia carlinoides
Banksia carlinoides, commonly known as Pink Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra carlinoides until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. polycephala (now B. polycephala
Banksia polycephala
Banksia polycephala, commonly known as Many-headed Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra polycephala until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. Kippistiana (now B. kippistiana
Banksia kippistiana
Banksia kippistiana is a shrub endemic to Western Australia. It was known as Dryandra kippistiana until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
Series Concinnæ
D. squarrosa (now B. squarrosa
Banksia squarrosa
Banksia squarrosa, commonly known as Pingle, is a shrub endemic to Western Australia.-Taxonomy:Specimens of B. squarrosa were first collected from near King George Sound in 1829 by William Baxter, and published by Robert Brown as Dryandra squarrosa the following year...

)
D. serra (now B. serra
Banksia serra
Banksia serra, commonly known as Serrate-leaved Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra polycephala until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.An assessment of the potential impact of climate change on this...

)
D. concinna (now B. concinna
Banksia concinna
Banksia concinna is a shrub endemic to Western Australia. It was known as Dryandra concinna until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. foliolata (now B. foliolata
Banksia foliolata
Banksia foliolata is a shrub endemic to Western Australia. It was known as Dryandra foliolata until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
Series Formosæ
D. stupposa (now B. stuposa
Banksia stuposa
Banksia stuposa is a shrub endemic to Western Australia. It was known as Dryandra stuposa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-Description:...

)
D. nobilis (now B. nobilis
Banksia nobilis
Banksia nobilis, commonly known as Golden Dryandra, Great Dryandra or Kerosene Bush, is a shrub endemic to Western Australia. It occurs on lateritic rises from Eneabba to Katanning in the state's Southwest Botanic Province. With large pinnatifid leaves with triangular lober, and a golden or reddish...

)
D. mucronulata (now B. mucronulata
Banksia mucronulata
Banksia mucronulata, commonly known as Swordfish Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra mucronulata until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. formosa (now B. formosa
Banksia formosa
Banksia formosa, commonly known as Showy Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra formosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. Baxteri (now B. biterax
Banksia biterax
Banksia biterax is a shrub endemic to Western Australia.It was known as Dryandra baxteri until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
Series Niveæ
Dryandra ser. Niveae
Dryandra ser. Niveae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into Banksia.-According to...

D. nivea (now B. nivea
Banksia nivea
Banksia nivea, commonly known as Honeypot Dryandra, is a shrub endemic to Western Australia. First described as Banksia nivea, it was transferred to Dryandra as Dryandra nivea by Robert Brown in 1810, and remained in that genus until 2007, when all Dryandra species were transferred to Banksia by...

)
D. arctotidis (now B. arctotidis
Banksia arctotidis
Banksia arctotidis is a shrub endemic to Western Australia. It was known as Dryandra arctotidis until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. arctotidis var. tortifolia (now B. tortifolia
Banksia tortifolia
Banksia tortifolia is a shrub endemic to Western Australia. It was first published as Dryandra tortifolia in 1855. In 1870, George Bentham demoted it to a variety of Dryandra arctotidis , but this was overturned by Alex George in 1999...

)
D. nana (now B. nana
Banksia nana
Banksia nana, commonly known as Dwarf Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra nana until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. Preissii (now B. acuminata
Banksia acuminata
Banksia acuminata is a rare prostrate shrub endemic to south-west Western Australia. It was published in 1848 as Dryandra preissii, but transferred into Banksia as B. acuminata in 2007.-Description:...

)
Series Obvallatæ
D. sclerophylla (now B. sclerophylla
Banksia sclerophylla
Banksia sclerophylla is a shrub endemic to Western Australia. It was known as Dryandra sclerophylla until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. pulchella (now B. bella
Banksia bella
Banksia bella, commonly known as the Silver Wongan Dryandra or Wongan Dryandra, is a shrub endemic to Western Australia. Unusual amongst dryandras, its foliage is glaucous-grey in colour....

)
D. plumosa (now B. plumosa
Banksia plumosa
Banksia plumosa is a shrub endemic to Western Australia. It was known as Dryandra plumosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. seneciifolia (now B. seneciifolia
Banksia seneciifolia
Banksia seneciifolia is a shrub endemic to Western Australia. It was known as Dryandra polycephala until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. vestita (now B. vestita
Banksia vestita
Banksia vestita, commonly known as Summer Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra vestita until 2007.-Description:...

)
D. cirsioides (now B. cirsioides
Banksia cirsioides
Banksia cirsioides is a shrub endemic to Western Australia. It was known as Dryandra cirsioides until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-Description:...

)
D. Hewardiana (now B. hewardiana
Banksia hewardiana
Banksia hewardiana is a shrub endemic to Western Australia. It was first published as Dryandra hewardiana by Carl Meissner in 1856. In 1870, George Bentham published what he held to be a closely related species under the name Dryandra patens, but in 1999 Alex George declared this a synonym of...

)
D. patens (now B. hewardiana)
D. conferta (now B. densa
Banksia densa
Banksia densa is a shrub endemic to Western Australia. It was known as Dryandra conferta until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. horrida (now B. horrida
Banksia horrida
Banksia horrida, commonly known as Prickly Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra horrida until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. serratuloides (now B. serratuloides
Banksia serratuloides
Banksia serratuloides is a shrub endemic to Western Australia. It was known as Dryandra serratuloides until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. comosa (now B. comosa
Banksia comosa
Banksia comosa, commonly known as Wongan Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra comosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
Series Gymnocephalæ
D. Shuttleworthiana (now B. shuttleworthiana
Banksia shuttleworthiana
Banksia shuttleworthiana, commonly known as Bearded Dryandra, is a shrub endemic to Western Australia, and is found between the towns of Geraldton and Gingin. It was known as Dryandra shuttleworthiana until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin...

)
D. speciosa (now B. splendida
Banksia splendida
Banksia splendida, commonly known as Shaggy Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra speciosa until 2007-Taxonomy:...

)
D. tridentata (now B. tridentata
Banksia tridentata
Banksia tridentata, commonly known as Yellow Honeypot, is a shrub endemic to Western Australia. It was published as Dryandra tridentata in 1856, and remained in that genus for 150 years, although in 1893 Benjamin Daydon Jackson accidentally listed it in Index Kewensis under Banksia, thus...

)
Section Aphragma
D. tenuifolia (now B. tenuis
Banksia tenuis
Banksia tenuis is a shrub endemic to Western Australia. It was known as Dryandra tenuifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...

)
D. proteoides (now B. proteoides
Banksia proteoides
Banksia proteoides, commonly known as King Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra proteoides until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. proteoides var. ferruginea (now B. rufa
Banksia rufa
Banksia rufa is a shrub endemic to Western Australia. It was first published by Carl Meisner in 1855, where it was given the name Dryandra ferruginea by Richard Kippist. The following year, Meisner published what was purportedly a distinct species, Dryandra runcinata...

D. runcinata (now B. r. subsp. rufa
Banksia rufa subsp. rufa
Banksia rufa subsp. rufa is a subspecies of Banksia rufa. It was known as Dryandra ferruginea subsp. ferruginea until 2007, when Austin Mast and Kevin Thiele sunk all Dryandra into Banksia. Since the name Banksia ferruginea had already been used, Mast and Thiele had to choose a new specific epithet...

)
D. obtusa (now B. obtusa
Banksia obtusa
Banksia obtusa, commonly known as Shining Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra obtusa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

)
D. bipinnatifida (now B. bipinnatifida
Banksia bipinnatifida
Banksia bipinnatifida is a shrub endemic to Western Australia.-Description:It grows as a prostrate shrub with an underground stem and a lignotuber. It has few leaves; these are deeply lobes, and the lobes are themselves deeply lobed, giving the impression of a bipinnate leaf structure...

)
D. pteridifolia (now B. pteridifolia
Banksia pteridifolia
Banksia pteridifolia, commonly known as Tangled Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra pteridifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...

)
D. calophylla (now B. calophylla
Banksia calophylla
Banksia calophylla is a shrub endemic to Western Australia. It was known as Dryandra calophylla until 2007, when all Dryandra species were transferred to Banksia by research botanists Austin Mast and Kevin Thiele.-External links:...

)

Legacy

Bentham's arrangement remained the accepted arrangement of Dryandra 126 years. It was finally replaced in 1996 by a new arrangement
George's taxonomic arrangement of Dryandra
Alex George's taxonomic arrangement of Dryandra was the first modern-day arrangement of that taxon. First published in Nuytsia in 1996, it superseded the arrangement of George Bentham, which had stood for over a hundred years; it would later form the basis for George's 1999 treatment of Dryandra...

 published by Alex George
Alex George
Alexander Segger George is a Western Australian botanist. He is the authority on the plant genera Banksia and Dryandra...

. George's arrangement retained a number of Bentham's infrageneric taxa.

In 2007, Austin Mast
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 an associate professor within the Department of Biological Science at Florida State University , and has been director of FSU's since August 2003.One of his...

 and Kevin Thiele
Kevin Thiele
Kevin R. Thiele is curator of the Western Australian Herbarium. His research interests include the systematics of the plant families Proteaceae, Rhamnaceae and Violaceae, and the conservation ecology of grassy woodland ecosystems...

 transferred Dryandra into Banksia, on the grounds that Banksia was paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...

with respect to Dryandra. Mast and Thiele were not yet ready to proffer an arrangement for the new circumscription of Banksia, so as an interim measure they transferred Dryandra at series rank, so as to minimise disruption to the nomenclature. Thus the infrageneric arrangement of Dryandra has been set aside, at least temporarily, and hence none of Bentham's infrageneric taxa are in current use.
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