Taxonomy of Banksia - AbsoluteAstronomy.com

Flowering plant

The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...

s, the taxonomy of Banksia is based on anatomical

Anatomy

Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

and morphological

Comparative anatomy

Comparative anatomy is the study of similarities and differences in the anatomy of organisms. It is closely related to evolutionary biology and phylogeny .-Description:...

properties of the Banksia

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. When it comes to size, banksias range from prostrate woody shrubs to trees up...

flower

Flower

A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

, fruiting structure

Fruit

In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

and seed

Seed

A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

, along with secondary characteristics such as leaf structure and growth habit. The genus is placed in family Proteaceae

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,...

, subfamily Grevilleoideae

Grevilleoideae

Grevilleoideae is a subfamily of the Proteaceae family of flowering plants. Mainly restricted to the southern hemisphere, it contains around 44 genera and about 950 species. Genera include Banksia, Grevillea and Macadamia.-Description:...

, tribe Banksieae and subtribe Banksiinae, alongside its close relative Dryandra

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 extensive molecular and morphological evidence that Banksia was paraphyletic with...

. The currently accepted infrageneric arrangement is based on Alex George

Alex George

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

's 1999 monograph for the Flora of Australia

Flora of Australia (series)

The Flora of Australia is a 59 volume series describing the vascular plants, bryophytes and lichens present in Australia and its external territories...

book series, and recognises two subgenera, three sections, 13 series, 77 species, 6 subspecies and 18 varieties. Recent cladistic analyses

Cladistics

Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

have cast doubt on aspects of this arrangement, but proposed alternative arrangements have not been widely accepted.

Background

Banksia is a genus

Genus

In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

of around 80 species

Species

In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

in the plant

Plant

Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

family Proteaceae

Proteaceae

. An iconic Australia

Australia

Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

n wildflower and popular garden

Garden

A garden is a planned space, usually outdoors, set aside for the display, cultivation, and enjoyment of plants and other forms of nature. The garden can incorporate both natural and man-made materials. The most common form today is known as a residential garden, but the term garden has...

plant

Plant

, they are easily recognised by their characteristic flower spikes and fruiting "cones". They grow in forms varying from prostrate woody shrub

Shrub

A shrub or bush is distinguished from a tree by its multiple stems and shorter height, usually under 5–6 m tall. A large number of plants may become either shrubs or trees, depending on the growing conditions they experience...

s to trees up to 35 metres tall, and occur in all but the most arid

Arid

A region is said to be arid when it is characterized by a severe lack of available water, to the extent of hindering or even preventing the growth and development of plant and animal life...

areas of Australia. As heavy producers of nectar

Nectar (plant)

Nectar is a sugar-rich liquid produced by plants. It is produced in glands called nectaries, either within the flowers, in which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide anti-herbivore protection...

, they are important sources of food for nectariferous animals such as honeyeater

Honeyeater

The honeyeaters are a large and diverse family of small to medium sized birds most common in Australia and New Guinea, but also found in New Zealand, the Pacific islands as far east as Samoa and Tonga, and the islands to the north and west of New Guinea known as Wallacea...

s and honey possum

Honey Possum

The honey possum or tait, its Native Australian name or noolbenger is a tiny Australian marsupial weighing just seven to eleven grams for the male, and eight to sixteen grams for the female—about half the weight of a mouse. Their physical size ranges from a body length of between 6.5 –...

s, and they are of economic importance to the nursery

Nursery (horticulture)

A nursery is a place where plants are propagated and grown to usable size. They include retail nurseries which sell to the general public, wholesale nurseries which sell only to businesses such as other nurseries and to commercial gardeners, and private nurseries which supply the needs of...

and cut flower

Floristry

Floristry is the general term used to describe production, commerce and trade in flowers. It encompasses flower care and handling, floral design or flower arranging, merchandising, and display and flower delivery. Wholesale florists sell bulk flowers and related supplies to professionals in the trade...

industries. However they are threatened by a number of processes, including land clearing, frequent burning, and disease; and a number of species are rare

Rare species

A rare species is a group of organisms that are very uncommon or scarce. This designation may be applied to either a plant or animal taxon, and may be distinct from the term "endangered" or "threatened species" but not "extinct"....

and endangered

Endangered species

An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

Taxonomic history

Specimens of Banksia were first collected by Sir Joseph Banks

Joseph Banks

Sir Joseph Banks, 1st Baronet, GCB, PRS was an English naturalist, botanist and patron of the natural sciences. He took part in Captain James Cook's first great voyage . Banks is credited with the introduction to the Western world of eucalyptus, acacia, mimosa and the genus named after him,...

and Dr Daniel Solander

Daniel Solander

Daniel Carlsson Solander or Daniel Charles Solander was a Swedish naturalist and an apostle of Carl Linnaeus. Solander was the first university educated scientist to set foot on Australian soil.-Biography:...

, naturalists on the Endeavour

HM Bark Endeavour

HMS Endeavour, also known as HM Bark Endeavour, was a British Royal Navy research vessel commanded by Lieutenant James Cook on his first voyage of discovery, to Australia and New Zealand from 1769 to 1771....

during Lieutenant (later Captain) James Cook

James Cook

Captain James Cook, FRS, RN was a British explorer, navigator and cartographer who ultimately rose to the rank of captain in the Royal Navy...

's first voyage to the Pacific Ocean

Pacific Ocean

The Pacific Ocean is the largest of the Earth's oceanic divisions. It extends from the Arctic in the north to the Southern Ocean in the south, bounded by Asia and Australia in the west, and the Americas in the east.At 165.2 million square kilometres in area, this largest division of the World...

. Cook landed on Australian soil for the first time on 29 April 1770, at a place that he later named Botany Bay

Botany Bay

Botany Bay is a bay in Sydney, New South Wales, a few kilometres south of the Sydney central business district. The Cooks River and the Georges River are the two major tributaries that flow into the bay...

in recognition of "the great quantity of plants Mr Banks and Dr Solander found in this place". 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: B. serrata

Banksia serrata

Banksia serrata, commonly known as Old Man Banksia, Saw Banksia, Saw-tooth Banksia and Red Honeysuckle, is a species of woody shrub or tree of the genus Banksia in the Proteaceae family. Native the east coast of Australia, it is found from Queensland through to Victoria with outlying populations on...

(Saw Banksia), B. integrifolia

Banksia integrifolia

Banksia integrifolia, commonly known as Coast Banksia, is a species of tree that grows along the east coast of Australia. One of the most widely distributed Banksia species, it occurs between Victoria and Central Queensland in a broad range of habitats, from coastal dunes to mountains...

(Coast Banksia), B. ericifolia

Banksia ericifolia

Banksia ericifolia, the Heath-leaved Banksia , is a species of woody shrub of the Proteaceae family native to Australia. It grows in two separate regions of Central and Northern New South Wales east of the Great Dividing Range...

(Heath-leaved Banksia) and B. robur

Banksia robur

Banksia robur, commonly known as Swamp Banksia or, less commonly, Broad-leaved Banksia grows in sand or peaty sand in coastal areas from Cooktown in north Queensland to the Illawarra region on the New South Wales south coast...

(Swamp Banksia). In June the ship was careened

Careening

Careening a sailing vessel is the practice of beaching it at high tide. This is usually done in order to expose one side or another of the ship's hull for maintenance and repairs below the water line when the tide goes out....

at Endeavour River

Endeavour River

The Endeavour River on Cape York Peninsula in Far North Queensland, Australia, was named in 1770 by Lt. James Cook, R.N., after he was forced to beach his ship, HM Bark Endeavour, for repairs in the river mouth, after damaging it on Endeavour Reef...

, where specimens of B. dentata

Banksia dentata

The Tropical Banksia is a species of small tree in the plant genus Banksia.-Distribution and habitat:It occurs throughout northern Australia, and also extends into New Guinea and the Aru Islands...

(Tropical Banksia) were collected.

Every specimen collected during the Endeavour voyage was sketched by Banks' botanical illustrator

Botanical illustrator

A botanical illustrator is a person who paints, sketches or otherwise illustrates botanical subjects such as trees and flowers. The job requires great artistic skill, attention to fine detail, and technical botanical knowledge...

Sydney Parkinson

Sydney Parkinson was a Scottish Quaker, botanical illustrator and natural history artist.Parkinson was employed by Joseph Banks to travel with him on James Cook's first voyage to the Pacific in 1768. Parkinson made nearly a thousand drawings of plants and animals collected by Banks and Daniel...

. On the Endeavour's return to England

England

England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

in July 1771, Banks' specimens became part of his London

London

London is the capital city of :England and the :United Kingdom, the largest metropolitan area in the United Kingdom, and the largest urban zone in the European Union by most measures. Located on the River Thames, London has been a major settlement for two millennia, its history going back to its...

herbarium

Herbarium

In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...

, and artists were employed to paint watercolours from Parkinson's sketches. Banks had plans to publish his entire collection as "Banks' Florilegium

Banks' Florilegium

Banks' Florilegium is a collection of copperplate engravings of plants collected by Sir Joseph Banks and Daniel Solander while they accompanied Captain James Cook on his voyage around the world between 1768 and 1771...

", but for various reasons the project was never completed, and it would be ten years before any of the Banksia species were formally published. By this time, a sixth species had been collected; in 1776, during Cook's third voyage, David Nelson

David Nelson (botanical collector)

David Nelson was gardener-botanist on the third voyage of James Cook, and botanist on the HMS Bounty under William Bligh at the time of the famous mutiny....

collected specimens of B. marginata

Banksia marginata

Banksia marginata, commonly known as the Silver Banksia, is a species of tree or woody shrub in the plant genus Banksia found throughout much of southeastern Australia. It ranges from the Eyre Peninsula in South Australia, to north of Armidale, New South Wales, and across Tasmania and the islands...

(Silver Banksia) from South Bruny Island, Tasmania

Tasmania

Tasmania is an Australian island and state. It is south of the continent, separated by Bass Strait. The state includes the island of Tasmania—the 26th largest island in the world—and the surrounding islands. The state has a population of 507,626 , of whom almost half reside in the greater Hobart...

.

The Banksia genus was finally described and named by Carolus Linnaeus the Younger

Carolus Linnaeus the Younger

Carl Linnaeus the Younger, Carl von Linné or Carolus Linnaeus the Younger was a Swedish naturalist...

in his April 1782 publication Supplementum Plantarum

Supplementum Plantarum

Supplementum Plantarum Systematis Vegetabilium Editionis Decimae Tertiae, Generum Plantarum Editiones Sextae, et Specierum Plantarum Editionis Secundae, commonly abbreviated to Supplementum Plantarum Systematis Vegetabilium or just Supplementum Plantarum, and further abbreviated by botanists to...

; hence the full name for the genus is "Banksia L.f.". Linnaeus placed the genus in class Tetrandra, order Monogynia of his father's classification, and named it in honour of Banks. The name Banksia had in fact already been published in 1775 as Banksia J.R.Forst

Johann Reinhold Forster

Johann Reinhold Forster was a German Lutheran pastor and naturalist of partial Scottish descent who made contributions to the early ornithology of Europe and North America...

& G.Forst

Georg Forster

Johann Georg Adam Forster was a German naturalist, ethnologist, travel writer, journalist, and revolutionary. At an early age, he accompanied his father on several scientific expeditions, including James Cook's second voyage to the Pacific...

, referring to some New Zealand

New Zealand

New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

species that the Forsters had collected during Cook's second voyage. However Linnaeus incorrectly attributed the Forsters' specimens to the genus Passerina

Passerina (plant)

Passerina is a genus in the plant family Thymelaeaceae.-List of species:*Passerina aragonensis*Passerina argentata*Passerina broteriana*Passerina burchellii*Passerina choulettei*Passerina clementii*Passerina comosa...

, and therefore considered the name Banksia available for use. By the time Joseph Gaertner

Joseph Gaertner

Joseph Gaertner was a German botanist, best known for his work on seeds, De Fructibus et Seminibus Plantarum ....

corrected Banks' error in 1788, Banksia L.f. was widely known and accepted, so Gaertner renamed

Nomen novum

In biological nomenclature, a nomen novum , new replacement name is a technical term...

Banksia J.R.Forst & G.Forst to Pimelia

Pimelia

Not to be confused with the plant genus Pimelea, which often is incorrectly spelt "Pimelia".Pimelia is a genus of darkling beetle in the subfamily Pimeliinae.There are many species, some of them listed at Wikispecies...

, a name previously chosen for the genus by Banks and Solander.

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

Costus

Costus is a genus of perennial tropical herbaceous plants from the costus family . They are often characterized and distinguished from relatives such as Zingiber by their spiraling stems. The genus as a whole is thus often called spiral gingers, but this can also refer to C...

L. In 1790 James Bruce

James Bruce

James Bruce was a Scottish traveller and travel writer who spent more than a dozen years in North Africa and Ethiopia, where he traced the origins of the Blue Nile.-Youth:...

published Bankesia Bruce, later corrected to Banksia Bruce, but the name was rejected in favour of Johann Friedrich Gmelin

Johann Friedrich Gmelin

Johann Friedrich Gmelin was a German naturalist, botanist, entomologist, herpetologist and malacologist.- Education :Johann Friedrich Gmelin was born as the eldest son of Philipp Friedrich Gmelin in 1748 in Tübingen...

's name Hagenia

Hagenia

Hagenia abyssinica is a species of flowering plant native to the high-elevation Afromontane regions of central and eastern Africa. It also has a disjunct distribution in the high mountains of East Africa from Sudan and Ethiopia in the north, through Kenya, Uganda, Rwanda, Burundi, Democratic...

. In 1820 the name Banksia Dombey

Joseph Dombey

Joseph Dombey was a French botanist. He was involved in the “Dombey affair” which was precipitated by British seizure of a vessel his collections were on and diversion of the collections to the British Museum.-Biography:He ran away from home and acquired a thorough knowledge of botany in...

ex DC.

A. P. de Candolle

Augustin Pyramus de Candolle also spelled Augustin Pyrame de Candolle was a Swiss botanist. René Louiche Desfontaines launched Candolle's botanical career by recommending him at an herbarium...

was published, but this was later determined by be a nomen nudum

Nomen nudum

The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...

that referred to the genus Cuphea

Cuphea

Cuphea is a genus containing about 260 species of annual and perennial flowering plants native to warm temperate to tropical regions of the Americas. The species range from low-growing herbs to semi-woody shrubs up to 2 m tall. Commonly they are known as cupheas, or, in the case of some species,...

, In 1891, Otto Kuntze

Otto Kuntze

Otto Carl Ernst Kuntze was a German botanist.-Biography:Otto Kuntze was born in Leipzig.An apothecary in his early career, he published an essay entitled Pocket Fauna of Leipzig. Between 1863 and...

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

James Britten

James Britten was an English botanist.-Biography:Born in Chelsea, London, he moved to High Wycombe in 1865 to begin a medical career. However he became increasingly interested in botany, and began writing papers on the subject...

's 1905 challenge. In 1940, Banksia L.f. was formally conserved against Banksia J.R.Forst. & G.Forst by Thomas Sprague.

In 1810, 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...

published descriptions and a taxonomic arrangement

Brown's taxonomic arrangement of Banksia

Robert Brown's taxonomic arrangement of Banksia was published in his 1810 Prodromus Florae Novae Hollandiae et Insulae Van Diemen, and expanded in this 1830 supplement to that publication, Supplementum Primum Prodromi Florae Novae Hollandiae...

of the 31 known species of Banksia in his Prodromus Florae Novae Hollandiae et Insulae Van Diemen

Prodromus Florae Novae Hollandiae et Insulae Van Diemen

Prodromus Florae Novae Hollandiae et Insulae Van Diemen is an 1810 flora of Australia by botanist Robert Brown. Often referred to as Prodromus Flora Novae Hollandiae, or by its standard botanical abbreviation Prodr. Fl. Nov. Holland., it was the first attempt at a survey of the Australian flora...

. He placed B. ilicifolia

Banksia ilicifolia

Banksia ilicifolia, commonly known as Holly-leaved Banksia, is a tree in the plant Proteaceae family. Endemic to southwest Western Australia, it belongs to Banksia subg. Isostylis, a subgenus of three closely related Banksia species with inflorescences that are dome-shaped heads rather than...

alone in subgenus Isostylis

Banksia subg. Isostylis

Banksia subg. Isostylis is a subgenus of Banksia. It contains three closely related species, all of which occur only in Southwest Western Australia. Members of subgenus Isostylis have dome-shaped flower heads that are superficially similar to those of B. ser...

, in recognition of its unusual dome-shaped inflorescence

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. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

. 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. Banksia verae was renamed Eubanksia by Stephan Endlicher

Stephan Ladislaus Endlicher

Stephan Ladislaus Endlicher was an Austrian botanist, numismatist and Sinologist. He was a director of the Botanical Garden of Vienna. He was born in Pressburg and died in Vienna....

in 1847.

By the time 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...

published his 1856 classification of the Proteaceae, there were 58 described Banksia species. Meissner's arrangement

Meissner's taxonomic arrangement of Banksia

Carl Meissner's taxonomic arrangement of Banksia was published in 1856, as part of his chapter on the Proteaceae in A. P. de Candolle's Prodromus systematis naturalis regni vegetabilis. It was the first attempt to provide an infrageneric classification for the genus, aside from Robert Brown's...

gave Isostylis and Eubanksia sectional rank, and divided the latter into four series based on leaf properties; these series were all highly heterogeneous.

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 :...

published his arrangement

Bentham's taxonomic arrangement of Banksia

George Bentham's taxonomic arrangement of Banksia was published in 1870, in Volume 5 of Bentham's Flora Australiensis. A substantial improvement on the previous arrangement, it would stand for over a century. It was eventually replaced by Alex George's 1981 arrangement, published in his classic...

of the Banksia in his landmark 1870 publication 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...

. 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

Pollen-presenter

A pollen-presenter is an area on the tip of the pistil in flowers of plants of the family Proteaceae. In this family, the anthers are difficult of access for potential pollination vectors such as bees, birds and nectariferous mammals....

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. Despite these shortcomings, this arrangement would stand for over 100 years.

Classification and relationships within Proteaceae

The framework for classification of genera within Proteaceae

Proteaceae

was laid by L. A. S. Johnson

Lawrence Alexander Sidney Johnson

Lawrence Alexander Sidney Johnson, known as Lawrie Johnson, was an Australian taxonomic botanist. He worked at the Royal Botanic Gardens, Sydney, for the whole of his professional career, as a botanist , Director and Honorary Research Associate .Alone or in collaboration with colleagues, he...

and Barbara Briggs

Barbara G. Briggs

Barbara Gillian Briggs is one of the foremost Australian botanists. The IK lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of the APG system....

in their influential 1975 monograph "On the Proteaceae: the evolution and classification of a southern family

On the Proteaceae: the evolution and classification of a southern family

On the Proteaceae: the evolution and classification of a southern family is a highly influential monograph on the evolution, biogeography and taxonomy of the flowering plant family Proteaceae. Authored by L. A. S...

". Their arrangement has been refined somewhat over the ensuing three decades, most notably by Peter Weston

Peter Weston

Peter Weston is a British science fiction fan. Now retired, he lives in Birmingham, UK.Weston's made many contributions in fan writing, fanzine editing, convention-running and in local science fiction clubs. His 1960s pseudonym "Malcolm Edwards" caused some confusion several years later, when a...

and Nigel Barker

Nigel Barker

Nigel Barker is an English author, spokesperson, filmmaker, fashion photographer and former model. He is best known for his participation as a judge and photographer on the reality show America's Next Top Model.-Early life:...

in 2006. Proteaceae is divided into five subfamilies, with Banksia placed in subfamily Grevilleoideae

Grevilleoideae

because the individual flowers in its inflorescence occur in pairs. On the basis of certain characters of the leaf venation, hairs and pollen

Pollen

Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

, it is grouped with three other genera in the tribe Banksieae. Two small genera, Austromuellera

Austromuellera

Austromuellera is a small genus of 2 species of rainforest tree from Northern Queensland. It was named in honour of Ferdinand von Mueller by Cyril Tenison White in 1930....

and Musgravea

Musgravea

Musgravea is a genus of rainforest tree from north-eastern Queensland.It was published in 1890 by Ferdinand von Mueller, and named in honour of Sir Anthony Musgrave, Governor of Queensland from 1883 to 1888. Together with its closest relative Austromuellera, it lies within the subtribe Musgraveinae...

, both of which occur only in the rainforest

Rainforest

Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

s of Queensland

Queensland

Queensland is a state of Australia, occupying the north-eastern section of the mainland continent. It is bordered by the Northern Territory, South Australia and New South Wales to the west, south-west and south respectively. To the east, Queensland is bordered by the Coral Sea and Pacific Ocean...

, are placed in subtribe Musgraveinae. Banksia is placed subtribe Banksiinae on a number of grounds of which the most obvious and easily recognised is the occurrence of flowers in condensed heads. The placement of Banksia in Proteaceae can be summarised as follows:

Family Proteaceae

Proteaceae

Subfamily Bellendenoideae

Subfamily Persoonioideae

Persoonioideae

The Persoonioideae are a subfamily of closely releted genera within the large and diverse Proteaceae family and incorporates such genera as Persoonia, Acidonia, Toronia and Placospermum....

Subfamily Symphionematoideae

Subfamily Proteoideae

Proteoideae

Proteoideae is one of five subfamilies of the flowering plant family Proteaceae. The greatest diversity of Proteoideae is in Africa, but there are also many species in Australia; a few species occur in South America, New Caledonia, and elsewhere.-Taxonomy:...

Subfamily Grevilleoideae

Grevilleoideae

Tribe Roupalae

Tribe Banksieae

Subtribe Musgraveinae

Subtribe Banksiinae

Genus Banksia

Banksia

Tribe Embothrieae

Tribe Macadamieae

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. Recent DNA

DNA

Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

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

Monophyly

In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

would be to sink Dryandra into Banksia.

Classical taxonomic treatment

In 1981, Alex George

Alex George

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

published his classic 1981 monograph The Genus Banksia L.f. (Proteaceae). George's arrangement was based on a variety of properties including leaf, style, pollen-presenter, follicle

Follicle (fruit)

In botany, a follicle is a dry unilocular many-seeded fruit formed from one carpel and dehiscing by the ventral suture in order to release seeds, such as in larkspur, magnolia, banksia, peony and milkweed....

and seed characters, with the criterion that a taxon was considered a distinct species only if it exhibited a "significant and consistent difference in the morphology of flowers and/or fruit". 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, which remains the standard text on the genus, and the treatment of Banksia in the Flora of Australia

Flora of Australia (series)

The Flora of Australia is a 59 volume series describing the vascular plants, bryophytes and lichens present in Australia and its external territories...

series.

George followed Brown in dividing Banksia into two subgenera, Banksia

Banksia 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 . Within this constraint, however, there have been various circumscriptions.-Banksia verae:B. subg...

and Isostylis. He then divided subgenus Banksia into two sections: Banksia

Banksia sect. Banksia

Banksia sect. Banksia is one of four sections of Banksia subgenus Banksia. It contains those species of subgenus Banksia with straight or sometimes curved but not hooked styles. These species all have cylindrical inflorescences and usually exhibit a bottom-up sequence of flower anthesis...

for species with straight or slightly curved styles, and Oncostylis

Banksia sect. Oncostylis

Banksia sect. Oncostylis is one of four sections of subgenus Banksia subg. Banksia. It contains those Banksia species with hooked pistils. All of the species in Oncostylis also exhibit a top-down sequence of flower anthesis, except for Banksia nutans which is bottom-up.Banksia sect...

for species with hooked styles. 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.

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

Banksia

Subgenus Banksia

Banksia subg. Banksia

Section Banksia

Banksia sect. Banksia

Series Salicinae

Banksia ser. Salicinae

Banksia ser. Salicinae is a valid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

B. dentata

Banksia dentata

The Tropical Banksia is a species of small tree in the plant genus Banksia.-Distribution and habitat:It occurs throughout northern Australia, and also extends into New Guinea and the Aru Islands...

– B. aquilonia

Banksia aquilonia

Banksia aquilonia is a species of tall shrub or tree in the plant genus Banksia.-Description:B. aquilonia grows as a tall shrub or small tree up to 8 metres high. It has hard, fissured, grey bark, and elliptic leaves without serrated margins...

– B. integrifolia

Banksia integrifolia

– B. plagiocarpa

Banksia plagiocarpa

Banksia plagiocarpa, commonly known as the Dallachy's banksia or blue banksia, is a species of shrub or tree in the plant genus Banksia. It occurs only on Hinchinbrook Island, Queensland and the immediately adjacent mainland....

– B. oblongifolia

Banksia oblongifolia

The Fern-leaved Banksia is a species of shrub in the plant genus Banksia. It occurs along the eastern coast of Australia from Wollongong, New South Wales in the south to Rockhampton, Queensland in the north...

– B. robur

Banksia robur

– B. conferta

Banksia conferta

The Glasshouse Banksia is a species of montane shrub or small tree in the plant genus Banksia. It occurs as distinct subspecies in two widely separated locations: B. conferta subsp. conferta in southern Queensland on the Lamington Plateau and further north in the Glass House Mountains; and...

– B. paludosa

Banksia paludosa

Banksia paludosa, commonly known as the marsh or swamp banksia, is a species of shrub in the plant genus Banksia. It is endemic to New South Wales, where it is found between Sydney and Batemans Bay, with an isolate population further south around Eden...

– B. marginata

Banksia marginata

– B. canei

Banksia canei

The mountain banksia is a species of shrub in the plant genus Banksia. It occurs in subalpine areas of the Great Dividing Range between Melbourne and Canberra in southeastern Australia...

– B. saxicola

Banksia saxicola

Banksia saxicola, commonly known as the Grampians Banksia is a species of tree or shrub in the plant genus Banksia. It occurs in Victoria in two distinct populations, one in The Grampians and the other on Wilsons Promontory....

Series Grandes

Banksia ser. Grandes

Banksia ser. Grandes is a taxonomic series in the genus Banksia. It consists of two closely related species in section Banksia, both endemic to Western Australia. These are B. grandis and B. solandri ....

B. grandis

Banksia grandis

Banksia grandis, commonly known as Bull Banksia, Giant Banksia or Mangite, is a common and distinctive tree in South West Western Australia....

– B. solandri

Banksia solandri

Banksia solandri, commonly known as Stirling Range Banksia, is a species of large shrub in the plant genus Banksia. It occurs only within the Stirling Range in southwest Western Australia.-Description:...

Series Banksia

Banksia ser. Banksia

Banksia ser. Banksia is avalid botanic name for a series of Banksia. As an autonym, it necessarily contains the type species of Banksia, B. serrata . Within this constraint, however, there have been various circumscriptions.-According to Bentham:Banksia ser. Banksia originated in 1870 as...

B. serrata

Banksia serrata

– B. aemula

Banksia aemula

Banksia aemula, commonly known as the wallum banksia, is a lignotuberous shrub of the Proteaceae family. Found from Bundaberg south to Sydney on the Australian east coast, it is encountered as a shrub or a taller tree to 8 m in coastal heath on deep sandy soil, known as Wallum...

– B. ornata

Banksia ornata

The Desert Banksia is a species of shrub in the plant genus Banksia which grows up to 3 m tall. It occurs in western Victoria, and in South Australia, where it is common on the Eyre Peninsula, Kangaroo Island and east of Adelaide, but is apparently absent from Yorke Peninsula.It tends to grow...

– B. baxteri

Banksia baxteri

The Baxter's Banksia , also known as Bird's Nest Banksia, is a species of shrub in the plant genus Banksia. It occurs along the south coast of Western Australia between Albany and Esperance....

– B. speciosa

Banksia speciosa

The Showy Banksia is a species of large shrub or small tree in the plant genus Banksia. It reaches up to 8 m in height...

– B. menziesii

Banksia menziesii

Banksia menziesii, commonly known as firewood banksia, is a species of flowering plant in the genus Banksia. It is a gnarled tree up to 10 m tall, or a lower spreading 1–3 m shrub in the more northern parts of its range. The serrated leaves are dull green with new growth a paler grey...

– B. candolleana

Banksia candolleana

The Propeller Banksia is a species of shrub in the plant genus Banksia. It occurs in sandplain country north of Gingin, Western Australia.-Description:...

– B. sceptrum

Banksia sceptrum

Banksia sceptrum, the Sceptre Banksia, occurs in Western Australia near the central west coast from Geraldton north through Kalbarri to Hamelin Pool. It extends inland almost to Mullewa....

Series Crocinae

Banksia ser. Crocinae

Banksia ser. Crocinae is a taxonomic series in the genus Banksia. It consists of four closely related species, all of which are endemic to Western Australia; namely B. prionotes , B. burdettii , B. hookeriana and B. victoriae...

B. prionotes

Banksia prionotes

Banksia prionotes, commonly known as Acorn Banksia or Orange Banksia, is a species of shrub or tree of the genus Banksia in the family Proteaceae. It is native to the southwest of Western Australia and can reach up to 10 m in height. It can be much smaller in more exposed areas or in the...

– B. burdettii

Banksia burdettii

Burdett's Banksia is a species of large shrub or small tree in the plant genus Banksia. It occurs on sandplain country north of Gingin, Western Australia....

– B. hookeriana

Banksia hookeriana

The Hooker's Banksia is a species of shrub in the plant genus Banksia. It occurs on sandplain shrubland between Arrowsmith and Eneabba in Western Australia. Seeds do not require any treatment, and take 23 to 49 days to germinate....

– B. victoriae

Banksia victoriae

Banksia victoriae, commonly known as Woolly Orange Banksia, is a species of large shrub or small tree in the plant genus Banksia...

Series Prostratae

Banksia ser. Prostratae

Banksia ser. Prostratae is a taxonomic series in the genus Banksia. It consists of six closely related species in section Banksia, all endemic to Western Australia, with a prostrate habit.Banksia ser...

B. goodii

Banksia goodii

Banksia goodii, commonly known as Good's Banksia, is an endangered shrub of Southwest Western Australia.Good's Banksia grows as a low shrub, either prostrate or with stems up to twenty centimetres high. The leaves are dark green with a prominent yellow midrib, and are held erect. They may be up...

– B. gardneri

Banksia gardneri

The Prostrate Banksia is a species of prostrate shrub in the plant genus Banksia. It occurs along the south coast of Western Australia.-Scientific name:...

– B. chamaephyton

Banksia chamaephyton

The Fishbone Banksia is a species of prostrate shrub in the plant genus Banksia. It occurs in sandplain country amongst low heath north of Perth, Western Australia.....

– B. blechnifolia

Banksia blechnifolia

Banksia blechnifolia is a species of prostrate shrub in the plant genus Banksia. Found in sandy soils in the south coastal region of Western Australia in the vicinity of Lake King, it is non-lignotuberous, regenerating by seed after bushfire...

– B. repens

Banksia repens

Banksia repens, the Creeping Banksia, is a species of shrub in the plant genus Banksia. It occurs on the south coast of Western Australia from D'Entrecasteaux National Park in the west to Mount Ragged in the east....

– B. petiolaris

Banksia petiolaris

Banksia petiolaris is a species of shrub of the genus Banksia in the Proteaceae family. It is a prostrate banksia, a group of several closely related species all with horizontal stems and thick, leathery upright leaves. No varieties have been subsequently described. It bears yellow inflorescences...

Series Cyrtostylis

Banksia ser. Cyrtostylis

Banksia ser. Cyrtostylis is a valid botanic name for a taxonomic series within the plant genus Banksia. First published at sectional rank by George Bentham in 1870, it was demoted to a series by Alex George in 1981. The name has had three circumscriptions.-According to Bentham:Bentham published B....

B. media

Banksia media

The Southern Plains Banksia , also known as Golden Stalk Banksia, is a species of shrub in the plant genus Banksia. It occurs on the south coast of Western Australia between Albany and Israelite Bay, where it is a common plant....

– B. praemorsa

Banksia praemorsa

The Cut-leaf Banksia is a species of shrub or tree in the plant genus Banksia. It occurs in a few isolated populations on the south coast of Western Australia between Albany and Cape Riche. Seeds do not require any treatment, and take 30 to 49 days to germinate.-References:...

– B. epica

Banksia epica

Banksia epica is a shrub that grows on the south coast of Western Australia. A spreading bush with wedge-shaped serrated leaves and large creamy-yellow flower spikes, it grows up to 3½ metres high. It is known only from two isolated populations in the remote south east of the state, near the...

– B. pilostylis

Banksia pilostylis

The Marsh Banksia is a species of shrub in the plant genus Banksia. It occurs on the south coast of Western Australia around Esperance. Seeds do not require any treatment, and take 18 to 49 days to germinate.-External links:...

– B. attenuata

Banksia attenuata

Banksia attenuata, commonly known as the candlestick banksia or slender banksia, is a species of plant in the proteaceae family. Commonly a tree, it reaches 10 m high, but is often a shrub in dryer areas 0.4 to 2 m high...

– B. ashbyi

Banksia ashbyi

The Ashby's Banksia is a species of shrub in the plant genus Banksia. It occurs in heath and spinifex country along the coast of Western Australia between Geraldton and Exmouth.-Description:...

– B. benthamiana

Banksia benthamiana

Bentham's Banksia is a species of shrub or tree in the plant genus Banksia. It occurs in scattered populations in the Wheatbelt region of Western Australia, one near Mullewa and the other near Dalwallinu.-Description:...

– B. audax

Banksia audax

Banksia audax is a species of shrub in the plant genus Banksia. It occurs over a large area in the central south of Western Australia.-Description:...

– B. lullfitzii

Banksia lullfitzii

Banksia lullfitzii is a species of shrub in the plant genus Banksia. A many-branched, spreading bush with golden-orange flowers, it occurs in scattered populations over a large area of the eastern goldfields of Western Australia...

– B. elderiana

Banksia elderiana

The Swordfish Banksia , commonly known as the swordfish banksia or palm banksia, is a species of shrub in the plant genus Banksia. It occurs in two disjunct areas in the Goldfields-Esperance region of Western Australia...

– B. laevigata

Banksia laevigata

Banksia laevigata is a species of shrub in the plant genus Banksia. It occurs in Western Australia's semi-arid shrubland from Southern Cross south to the Fitzgerald River National Park. It is composed of two closely related subspecies, B. laevigata subsp. laevigata and B. laevigata...

– B. elegans

Banksia elegans

The Elegant Banksia is a species of shrub or small tree in the plant genus Banksia. It occurs only over a 65 square kilometre area north and west of Eneabba, Western Australia.-Ecology:...

– B. lindleyana

Banksia lindleyana

Banksia lindleyana, commonly known as the Porcupine Banksia, is a species of woody shrub in the genus Banksia of the family Proteaceae. It generally grows as a small shrub to 1 m high with long narrow serrated leaves, and bright yellow oval or round inflorescences...

Series Tetragonae

Banksia ser. Tetragonae

Banksia ser. Tetragonae is a taxonomic series in the genus Banksia. It consists of three closely related species of erect shrub with pendulous inflorescences in section Banksia. These are B. lemanniana , B. caleyi and B. aculeata ....

B. lemanniana

Banksia lemanniana

Banksia lemanniana, commonly known as the Yellow lantern Banksia or Lemann's Banksia, is a species of woody shrub in the genus Banksia of the family Proteaceae. It generally grows as an open shrub or small tree to 5 m high with stiff serrated leaves, and unusually hanging inflorescences...

– B. caleyi

Banksia caleyi

Banksia caleyi, commonly known as the red lantern banksia or Cayley's banksia, is a species of shrub in the plant genus Banksia. It occurs in southern Western Australia south and east of the Stirling Ranges through to around Jerramungup...

– B. aculeata

Banksia aculeata

Banksia aculeata, commonly known as Prickly Banksia, is a species of shrub native to the Stirling Range in southwest Western Australia. One of three closely related species all of which have distinctive upside down lantern-like inflorescences, it bears pinkish blooms in early summer. The leaves...

Series Bauerinae

B. baueri

Banksia baueri

The Woolly Banksia is a species of shrub in the plant genus Banksia. It occurs in southwest Western Australia north and east of Albany. It has a distinctively large and hairy looking inflorescence which can be 300 mm or more long and up to 200 mm in diameter.It is placed alone in series...

Series Quercinae

Banksia ser. Quercinae

Banksia ser. Quercinae is avalid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

B. quercifolia

Banksia quercifolia

The Oak-leaved Banksia is a species of shrub in the plant genus Banksia. It occurs on the south coast of Western Australia from D'Entrecasteaux National Park in the west to Mount Manypeaks in the east.-Ecology:...

– B. oreophila

Banksia oreophila

The Western Mountain Banksia or Mountain Banksia is a species of shrub in the plant genus Banksia. It occurs on the slopes and hilltops of the Stirling and Barren Ranges in southwest Western Australia.-Taxonomy:...

Section Coccinea

B. coccinea

Banksia coccinea

Banksia coccinea, commonly known as the Scarlet Banksia, Waratah Banksia or Albany Banksia, is an erect shrub or small tree in the plant genus Banksia...

Section Oncostylis

Banksia sect. Oncostylis

Series Spicigerae

Banksia ser. Spicigerae

Banksia ser. Spicigerae is a taxonomic series in the genus Banksia. It consists of the seven species in section Oncostylis that have cylindrical inflorescences. These range in form from small shrubs to tall trees. The leaves grow in either an alternate or whorled pattern, with various shape forms...

B. spinulosa

Banksia spinulosa

The Hairpin Banksia is a species of woody shrub, of the genus Banksia in the Proteaceae family, native to eastern Australia. Widely distributed, it is found as an understorey plant in open dry forest or heathland from Victoria to northern Queensland, generally on sandstone though sometimes also...

– B. ericifolia

Banksia ericifolia

– B. verticillata

Banksia verticillata

Banksia verticillata, commonly known as Granite Banksia or Albany Banksia, is a species of shrub or tree of the genus Banksia in the Proteaceae family. It is native to the southwest of Western Australia and can reach up to 3 m in height. It can grow taller to 5 m in sheltered areas,...

– B. seminuda

Banksia seminuda

Banksia seminuda, commonly known as the River Banksia, is a tree in the plant genus Banksia. It is found in south west Western Australia from Dwellingup to the Broke Inlet east of Denmark . It is often mistaken for and was originally considered a subspecies of the Banksia littoralis...

– B. littoralis

Banksia littoralis

Banksia littoralis, commonly known as the Swamp Banksia, Swamp Oak, Pungura and the Western Swamp Banksia, is a tree in the plant genus Banksia. It is found in south west Western Australia from the south eastern metropolitan area of Perth to the Stirling Range and Albany...

– B. occidentalis

Banksia occidentalis

The Red Swamp Banksia or Waterbush is a species of shrub or small tree in the plant genus Banksia. It occurs on the south coast of Western Australia in three disjunct populations: at Augusta, around Albany and in the Esperance area.A 1980 field study at Cheyne beach showed it to be pollinated by...

– B. brownii

Banksia brownii

Banksia brownii, commonly known as Feather-leaved Banksia or Brown's Banksia, is a species of shrub that occurs in southwest Western Australia. An attractive plant with fine feathery leaves and large red-brown flower spikes, it usually grows as an upright bush around two metres high, but can also...

Series Tricuspidae

B. tricuspis

Banksia tricuspis

The Lesueur Banksia or Pine Banksia is a species of shrub or tree in the plant genus Banksia. It occurs within a geographic range of just 15 square kilometres near Jurien, Western Australia.-External links:...

Series Dryandroidae

B. dryandroides

Banksia dryandroides

Banksia dryandroides, the Dryandra-leaved Banksia, is a species of small shrub in the plant genus Banksia. It occurs in shrubland, coastal heath and woodland on the south coast of Western Australia between Two Peoples Bay and Cheyne Bay. The species is placed alone in series B. ser...

Series Abietinae

Banksia ser. Abietinae

Banksia ser. Abietinae is avalid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

B. sphaerocarpa

Banksia sphaerocarpa

Banksia sphaerocarpa, commonly known as the Fox Banksia or Round-fruit Banksia, is a species of shrub or tree in the plant genus Banksia . It is generally encountered as a 1–2 m high shrub, and is usually smaller in the north of its range...

– B. micrantha

Banksia micrantha

Banksia micrantha is a species of shrub in the plant genus Banksia. A small spreading bush with pale yellow flower spikes, it occurs between Eneabba and Cervantes in South west Western Australia...

– B. grossa

Banksia grossa

Banksia grossa, commonly known as Coarse Banksia is a species of shrub in the plant family Proteaceae endemic to south west Western Australia. One of fourteen species of banksia with predominantly round or oval inflorescences of the series Abietinae, it was described in 1981 as a distinct species...

– B. telmatiaea

Banksia telmatiaea

Banksia telmatiaea, commonly known as Swamp Fox Banksia or rarely Marsh Banksia, is a shrub that grows in marshes and swamps along the lower west coast of Australia. It grows as an upright bush up to 2 m tall, with narrow leaves and a pale brown flower spike, which can produce profuse...

– B. leptophylla

Banksia leptophylla

The Slender-leaved Banksia is a species of shrub in the plant genus Banksia. It occurs along the west coast of Western Australia from Gingin to Kalbarri. Before Alex George's revision of 1981, it was labelled informally as B. sphaerocarpa var. pinifolia or var...

– B. lanata

Banksia lanata

The Coomallo Banksia is a species of shrub in the plant genus Banksia. It occurs within a range of less than 100 square kilometres between Eneabba and Mount Lesueur, Western Australia. It has roughly spherical inflorescences with flowers of cream to orange-brown colour. The leaves are linear and...

– B. scabrella

Banksia scabrella

Banksia scabrella, commonly known as the Burma Road Banksia, is a species of woody shrub in the genus Banksia. It is classified in the series Abietinae, a group of several species of shrubs with small round or oval inflorescences...

– B. violacea

Banksia violacea

Banksia violacea, commonly known as Violet Banksia, is a species of shrub or tree in the plant genus Banksia . It generally grows as a small shrub to 1.5 m high with fine narrow leaves, and is best known for its unusually coloured dark purple-violet inflorescences...

– B. incana

Banksia incana

The Hoary Banksia is a species of small shrub in the plant genus Banksia. It occurs on sandplain heathland between Badgingarra and Eneabba in Western Australia, with outlying populations as far south as Perth. Seeds do not require any treatment, and take around 14 days to germinate....

– B. laricina

Banksia laricina

The Rose-Fruited Banksia is a species of shrub in the plant genus Banksia endemic to southwestern Western Australia. It derives its specific Latin name from larix or larch, which its foliage is said to resemble. The common name comes from the striking fruits which resemble wooden roses...

– B. pulchella

Banksia pulchella

The Teasel Banksia is a species of small shrub in the plant genus Banksia. It occurs on the south coast of Western Australia from Fitzgerald River National Park east to Israelite Bay....

– B. meisneri

Banksia meisneri

The Meisner's Banksia is a species of small shrub in the plant genus Banksia. It occurs in a number of isolated populations throughout southwest Western Australia. Seeds do not require any treatment, and take 28 to 39 days to germinate.-External links:...

– B. nutans

Banksia nutans

Banksia nutans, commonly known as Nodding Banksia, is a species of shrub native to the south coast of Western Australia in the genus Banksia...

Subgenus Isostylis

Banksia subg. Isostylis

B. ilicifolia

Banksia ilicifolia

– B. oligantha

Banksia oligantha

Banksia oligantha, commonly known as Wagin Banksia, is an endangered species in the plant family Proteaceae endemic to south west Western Australia. It belongs to Banksia subg. Isostylis, a subgenus of three closely related Banksia species with dome-shaped heads as inflorescences, rather than...

– B. cuneata

Banksia cuneata

Banksia cuneata, commonly known as Matchstick Banksia or Quairading Banksia, is an endangered species of flowering plants in the Proteaceae family. Endemic to southwest Western Australia, it belongs to Banksia subg...

Thiele and Ladiges

In 1996, 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...

and Pauline Ladiges published a cladistic analysis of the Banksia genus in the journal Australian Systematic Botany

Australian Systematic Botany

Australian Systematic Botany is a peer-reviewed international journal that publishes original research, and sometimes review articles, on topics related to systematic botany, such as biogeography, taxonomy and evolution. It is published by CSIRO Publishing...

. As their cladogram

Cladistics

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 B. conferta subsp. penicillata

Banksia conferta subsp. penicillata

The Newnes Plateau Banksia is a plant only described in 1981 though collected in the 1970s...

); B. gardneri var. brevidentata

Banksia gardneri var. brevidentata

Banksia gardneri var. brevidentata is a variety of Banksia gardneri. It is native to the Southwest Botanical Province of Western Australia....

to B. brevidentata; B. gardneri var. hiemalis

Banksia gardneri var. hiemalis

Banksia gardneri var. hiemalis is a variety of Banksia gardneri. It is native to the Southwest Botanical Province of Western Australia. Seeds do not require any treatment, and take around 19 days to germinate....

to B. hiemalis; and B. sphaerocarpa var. dolichostyla

Banksia sphaerocarpa var. dolichostyla

Banksia sphaerocarpa var. dolichostyla, commonly known as Ironcap Banksia, is a plant which is either considered a variety of Banksia sphaerocarpa, or as a species in its own right. It is native to the Southwest Botanical Province of Western Australia...

to B. dolichostyla. Two new series and eleven subseries 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

Incertae sedis

, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

.

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. However, a number of Australian herbaria

Herbarium

have continued to follow Thiele and Ladiges on some points, for example by recognising the four species that they promoted.

DNA analysis

Following on from an earlier molecular study, 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 co-authors published cladistic analyses of genetic data from DNA

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

Stoma

In botany, a stoma is a pore, found in the leaf and stem epidermis that is used forgas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that are responsible for regulating the size of the opening...

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 B. attenuata

Banksia attenuata

.

Thus, the results presented by Mast et al. strongly suggest that Banksia is paraphyletic with respect to 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.

The change has been adopted by the Western Australian Herbarium, now headed by Thiele, but Banksia authority Alex George has strongly challenged it.

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