Banksia menziesii
Encyclopedia
Banksia menziesii, commonly known as firewood banksia, is a 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...

 of flowering plant in the genus 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...

. It is a gnarled tree up to 10 m (35 ft) tall, or a lower spreading 1–3 m (4–10 ft) shrub in the more northern parts of its range. The serrated leaves are dull green with new growth a paler grey green. The prominent autumn and winter 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...

s are often two-coloured red or pink and yellow, and their colour has given rise to more unusual common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

s such as port wine banksia and strawberry banksia. Yellow blooms are rarely seen.

First described by the botanist 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 the early 19th century, no separate varieties of Banksia menziesii are recognised. It is found in 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...

, from the Perth
Perth, Western Australia
Perth is the capital and largest city of the Australian state of Western Australia and the fourth most populous city in Australia. The Perth metropolitan area has an estimated population of almost 1,700,000....

 (32° S) region north to the Murchison River
Murchison River (Western Australia)
The Murchison River is the second longest river in Western Australia. It flows for about from the southern edge of the Robinson Ranges to the Indian Ocean at Kalbarri. It has a mean annual flow of about 200 million cubic metres.-Course:...

 (27° S), and generally grows on sandy soils, in scrubland or low woodland
Woodland
Ecologically, a woodland is a low-density forest forming open habitats with plenty of sunlight and limited shade. Woodlands may support an understory of shrubs and herbaceous plants including grasses. Woodland may form a transition to shrubland under drier conditions or during early stages of...

. Banksia menziesii provides food for a wide array of invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

 and vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

 animals; birds and in particular 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 are prominent visitors. A relatively hardy plant, Banksia menziesii is commonly seen in gardens, nature strips
Tree lawn
A road verge, is a narrow strip of grass or plants and sometimes also trees typically located beside the carriageway within the boundary of a road.The land is often public...

 and parks in Australian urban areas with Mediterranean climate
Mediterranean climate
A Mediterranean climate is the climate typical of most of the lands in the Mediterranean Basin, and is a particular variety of subtropical climate...

s, but its sensitivity to dieback from the soil-borne water mould Phytophthora cinnamomi
Phytophthora cinnamomi
Phytophthora cinnamomi is a soil-borne water mould that produces an infection which causes a condition in plants called root rot or dieback. The plant pathogen is one of the world's most invasive species and is present in over 70 countries from around the world.- Life cycle and effects on plants :P...

 makes it short-lived in places with humid summers, such as Sydney
Sydney
Sydney is the most populous city in Australia and the state capital of New South Wales. Sydney is located on Australia's south-east coast of the Tasman Sea. As of June 2010, the greater metropolitan area had an approximate population of 4.6 million people...

. Banksia menziesii is widely used in the cut flower industry
Floriculture
Floriculture, or flower farming, is a discipline of horticulture concerned with the cultivation of flowering and ornamental plants for gardens and for floristry, comprising the floral industry...

 both in Australia and overseas.

Description

Banksia menziesii grows either as a gnarled tree to 10 m (35 ft), or a lower spreading 1–3 metre (4–10 ft) shrub, generally encountered at its northern limits in the vicinity of Eneabba
Eneabba, Western Australia
Eneabba is a town on the Brand Highway located 278 km north of Perth, Western Australia.The area is famous for its spectacular display of wildflowers in the spring. It is also home to the Iluka Resources mineral sands facility....

-Mount Adams; thus, it declines steadily in size as the climate becomes warmer and dryer further north. In the shrub form, several stems arise from the woody base known as the lignotuber
Lignotuber
A lignotuber is a starchy swelling of the root crown possessed by some plants as a protection against destruction of the plant stem by fire. The crown contains buds from which new stems may sprout, and a sufficient store of nutrients to support a period of growth in the absence of...

. The trunk is greyish, sometimes with shades of brown or pink, and the 2–3 cm (1 in) thick rough bark breaks away easily. The new growth is covered in fine brownish hair, which wears away after two or three years, leaving smooth stems and leaves. Stems which will bear flower spikes the following year are generally thicker and longer. Oblong in shape and somewhat truncate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...

 at the tips, the leaves are grey-green in colour, 8–25 cm (3–10 in) long and up to 4 cm (1.6 in) wide. The new leaves are paler and finely downy. The leaf margins are serrated with many small 1–2 mm long triangular teeth. The lower surface of the leaf has a midrib covered in fine pale brown hair.

Flowering occurs in autumn and winter, peaking from May to July. Overall the 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...

s, or flower spikes, take around eight months to development from the first microscopic changes in late spring. Ovoid to cylindrical in shape, the flower spikes can be up to 7–8 cm (2.6–3.4 in) wide and 4–12 cm (1.6–4.8 in) high. They are composed of numerous individual flowers; one field study south of Perth recorded an average of 1043 per flower spike, while another on plants in cultivation in South Australia recorded an average of 720. B. menziesii has more flower colour variants than any other Banksia species, with flower spikes occurring in a wide range of pinks, as well as chocolate, bronze, yellow and white, and greenish variants. They are particularly striking closeup but can look indistinct from a distance. They are most attractive in late bud, the styles contrasting well to the body of the inflorescence, the whole looking like a red- or pink-and white vertical candy striped bloom. The inflorescences are generally a deeper red after colder weather and further into the winter. Anthocyanin
Anthocyanin
Anthocyanins are water-soluble vacuolar pigments that may appear red, purple, or blue according to pH...

 pigments are responsible for the red and pink shades in the flowers.

Old flowers usually fall off the spikes quickly, with up to 25 large beaked follicles developing. A mottle
Mottle
Mottle or mottling is the appearance of uneven spots. It is commonly used to describe plants or the skin of animals. In plants, mottling usually consists of yellowish spots on plants, and is usually a sign of disease or malnutrition...

d dark brown and grey in colour, these can be prominent and quite attractively patterned when newly developed. Oval shaped, they are 2.5–3.5 cm (1–1.4 in) long by 1–1.5 cm (0.4–0.6 in) high and 1–1.5 cm (0.4–0.6 in) wide. Overall, only a small fraction of flowers develop into follicles; the proportion is as low as one in a thousand. The plant is dependent on fire to reproduce as the follicles only open after being burnt, each one producing one or two viable wedge-shaped (cuneate) seeds, on either side of a woody separator. The colour and level of pigmentation in the seeds foreshadows the eventual colour of the inflorescences. Kevin Collins of the Banksia Farm recalled that for many years pale seeds were discarded by seed collectors who thought they were infertile. Later, he learnt that pale seeds yielded yellow-coloured blooms, dark grey the usual red-coloured, and black a distinctive bronze-coloured bloom.

Seedlings have obovate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...

 cotyledons 1–1.4 cm (0.4–0.6 in) long by 1–1.5 cm (0.4–0.6 in) wide, and the leaves which develop immediately afterward are crowded and very hairy. They have serrate margins. Evidence of thickening to form a future lignotuber, as well as minute buds, has been detected from the bases of seedlings at five months of age.

Taxonomy

The common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 of firewood banksia was a result of its quick-burning properties. Other names recorded include Menzies banksia, firewheel banksia, port wine banksia, flame banksia, and in the cut flower
Floriculture
Floriculture, or flower farming, is a discipline of horticulture concerned with the cultivation of flowering and ornamental plants for gardens and for floristry, comprising the floral industry...

 industry, strawberry banksia and raspberry frost banksia. The Beeloo Whadjuk
Whadjuk
Whadjuk, also called Wadjuk, Whajook and Wadjug, is the name according to Norman Tindale for the Aboriginal group inhabiting the Western Australian region of the Perth bioregion of the Swan Coastal Plain, and extending below Walyunga into the surrounding Jarrah Forests...

 Noongar
Noongar
The Noongar are an indigenous Australian people who live in the south-west corner of Western Australia from Geraldton on the west coast to Esperance on the south coast...

 people of the Perth region knew it as the Mungyt. Despite its variation across its range, George noted that B. menziesii was a clearly defined species, and no formal division into subspecies was warranted.

History

Specimens of B. menziesii were first collected by the botanist Charles Fraser
Charles Fraser (botanist)
Charles Fraser or Frazer was Colonial Botanist of New South Wales from 1821 to 1831. He collected and catalogued numerous Australian plant species, and participated in a number of exploring expeditions...

 during Captain (later Admiral Sir) James Stirling
James Stirling (Australian governor)
Admiral Sir James Stirling RN was a British naval officer and colonial administrator. His enthusiasm and persistence persuaded the British Government to establish the Swan River Colony and he became the first Governor and Commander-in-Chief of Western Australia...

's March 1827 exploration of the Swan River
Swan River (Western Australia)
The Swan River estuary flows through the city of Perth, in the south west of Western Australia. Its lower reaches are relatively wide and deep, with few constrictions, while the upper reaches are usually quite narrow and shallow....

. The following year, Alexander Macleay
Alexander Macleay
Hon. Alexander Macleay MLC FLS FRS was a leading member of the Linnean Society and a fellow of the Royal Society.Macleay was born on Ross-shire, Scotland, eldest son of William Macleay, provost of Wick...

 sent some of Fraser's specimens to 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...

. Brown formally published the species in his 1830 Supplementum Primum Prodromi Florae Novae Hollandiae
Supplementum Primum Prodromi Florae Novae Hollandiae
Supplementum primum Prodromi florae Novae Hollandiae is an 1830 supplement to Robert Brown's Prodromus florae Novae Hollandiae et Insulae Van Diemen. It may be referred to by its standard botanical abbreviation Suppl. Prodr. Fl. Nov...

, giving it the specific epithet in honor of Archibald Menzies
Archibald Menzies
Archibald Menzies was a Scottish surgeon, botanist and naturalist.- Life and career :Menzies was born at Easter Stix in the parish of Weem, in Perthshire. While working with his elder brother William at the Royal Botanic Gardens, he drew the attention of Dr John Hope, professor of botany at...

, surgeon-naturalist on the HMS Discovery
HMS Discovery (1789)
HMS Discovery was a Royal Navy ship launched in 1789 and best known as the lead ship in George Vancouver's exploration of the west coast of North America in his famous 1791-1795 expedition. She was converted to a bomb vessel in 1798 and participated in the Battle of Copenhagen. Thereafter she...

 under George Vancouver
George Vancouver
Captain George Vancouver RN was an English officer of the British Royal Navy, best known for his 1791-95 expedition, which explored and charted North America's northwestern Pacific Coast regions, including the coasts of contemporary Alaska, British Columbia, Washington and Oregon...

, who discovered King George Sound
King George Sound
King George Sound is the name of a sound on the south coast of Western Australia. Located at , it is the site of the city of Albany.The sound covers an area of and varies in depth from to ....

 in 1791. Thus the species' full name is Banksia menziesii R.Br. Neither Brown nor Menzies ever saw the plant growing.

Under Brown's 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...

, B. menziesii was placed in subgenus
Subgenus
In biology, a subgenus is a taxonomic rank directly below genus.In zoology, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the generic name and the specific epithet: e.g. the Tiger Cowry of the Indo-Pacific, Cypraea tigris Linnaeus, which...

 Banksia verae, the "True Banksias", because its inflorescence is a typical Banksia flower spike. 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, and demoted to sectional
Section (botany)
In botany, a section is a taxonomic rank below the genus, but above the species. The subgenus, if present, is higher than the section, and the rank of series, if present, is below the section. Sections are typically used to help organise very large genera, which may have hundreds of species...

 rank by 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...

 in his 1856 classification
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...

. Meissner further divided Eubanksia into four series
Series (botany)
Series is a low-level taxonomic rank below that of section but above that of species.In botany, a series is a subdivision of a genus...

, with B. menziesii placed in 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:...

. When 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 1870 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...

 in 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 discarded Meissner's series, replacing them with four sections
Section (botany)
In botany, a section is a taxonomic rank below the genus, but above the species. The subgenus, if present, is higher than the section, and the rank of series, if present, is below the section. Sections are typically used to help organise very large genera, which may have hundreds of species...

. B. menziesii was placed in Orthostylis, a somewhat heterogeneous section containing 18 species. This arrangement would stand for over a century.

In 1891, German botanist 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...

 challenged the generic name Banksia L.f.
Carolus Linnaeus the Younger
Carl Linnaeus the Younger, Carl von Linné or Carolus Linnaeus the Younger was a Swedish naturalist...

, on the grounds that the name Banksia had previously 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 the genus now known as Pimelea
Pimelea
Pimelea is a genus of plants belonging to the family Thymelaeaceae. There are about 80 species in the genus, native to Australia and New Zealand. Many of the species are poisonous to cattle.Selected species...

. Kuntze proposed Sirmuellera as an alternative, republishing B. menziesii as "Sirmuellera menziesii". The challenge failed, Banksia L.f. was formally conserved, and Sirmuellera menziesii (R.Br.) Kuntze is now a nomenclatural synonym of B. menziesii.

Current placement

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

 published a new taxonomic arrangement of Banksia in his classic 1981 monograph The genus Banksia L.f. (Proteaceae)
The genus Banksia L.f. (Proteaceae)
The genus Banksia L.f. is a 1981 monograph by Alex George on the taxonomy of the plant genus Banksia. Published by the Western Australian Herbarium as Nuytsia 3, it presented George's taxonomic arrangement of Banksia, the first major taxonomic revision of the genus since George Bentham published...

. Endlicher's Eubanksia became B. subg. 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 was divided into three sections. B. menziesii was placed in B. sect. 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...

, and this was further divided into nine series, with B. menziesii placed in B. ser. 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...

.
He thought its closest relatives to be Banksia speciosa and B. baxteri, and that it also formed a link with eastern species, particularly Banksia serrata. Since Brown's original publication had treated all of Fraser's specimens as syntype
Syntype
In biological nomenclature, a syntype is a term used to indicate a specimen with a special status.In zoological nomenclature, a syntype is defined as "Each specimen of a type series from which neither a holotype nor a lectotype has been designated [Arts. 72.1.2, 73.2, 74]. The syntypes...

s for the species, George also chose a lectotype
Lectotype
In botanical nomenclature and zoological nomenclature, a lectotype is a kind of name-bearing type. When a species was originally described on the basis of a name-bearing type consisting of multiple specimens, one of those may be designated as the lectotype...

, selecting a tree that Fraser had received from Macleay in May 1828.

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 new arrangement for the genus, after cladistic
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...

 analyses yielded a cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...

 significantly different from George's arrangement. Thiele and Ladiges' arrangement
Thiele and Ladiges' taxonomic arrangement of Banksia
Kevin Thiele and Pauline Ladiges' taxonomic arrangement of Banksia, published in 1996, was a novel taxonomic arrangement that was intended to align the taxonomy of Banksia more closely with the phylogeny that they had inferred from their cladistic analysis of the genus...

 retained B. menziesii in series Banksia, placing it in B. subser. Cratistylis
Banksia subser. Cratistylis
Banksia subser. Cratistylis is a valid botanic name for a subseries of Banksia. It was first published by Kevin Thiele in 1996, but discarded by Alex George in 1999.-Cladistics:...

 along with nine other species. This arrangement stood until 1999, when George effectively reverted to his 1981 arrangement in his 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...

 series.

Under George's taxonomic arrangement of Banksia
George's taxonomic arrangement of Banksia
Alex George's taxonomic arrangement of Banksia was the first modern-day arrangement for that genus. First published in 1981 in the classic monograph The genus Banksia L.f. , it superseded the arrangement of George Bentham, which had stood for over a hundred years. It was overturned in 1996 by Kevin...

, B. menziesii's taxonomic placement may be summarised as follows:
Genus 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...

Subgenus 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...

Section 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...

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
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...

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
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....



B. menziesii's inflorescences resemble those of no other banksia; its closest relatives were felt by George to be B. speciosa (showy banksia) and B. baxteri (bird's nest banksia), which differs from B. menziesii in having yellow flowers and leaves with deep triangular lobes. In 2002, a molecular study by 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...

 showed its closest relatives to be the members of the 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...

.

In 2005, Mast, Eric Jones and Shawn Havery published the results of their cladistic analyses of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 data for Banksia. They inferred a phylogeny greatly different from the accepted taxonomic arrangement, including finding Banksia to be 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. 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
Banksia subg. Spathulatae
Banksia subg. Spathulatae is a valid botanic name for a subgenus of Banksia. It was published in 2007 by Austin Mast and Kevin Thiele, and defined as containing all those Banksia species having spathulate cotyledons...

 for the species having spoon-shaped cotyledon
Cotyledon
A cotyledon , is a significant part of the embryo within the seed of a plant. Upon germination, the cotyledon may become the embryonic first leaves of a seedling. The number of cotyledons present is one characteristic used by botanists to classify the flowering plants...

s; in this way they also redefined the autonym
Autonym (botany)
In botanical nomenclature, autonyms are automatically created names, as regulated by the International Code of Botanical Nomenclature . Autonyms are cited without an author. Relevant provisions are in articles 6.8, 22.1-3 and 26.1-3....

 B. subg. 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...

. They foreshadowed publishing a full arrangement once DNA sampling
Sampling (statistics)
In statistics and survey methodology, sampling is concerned with the selection of a subset of individuals from within a population to estimate characteristics of the whole population....

 of Dryandra was complete. In the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then B. menziesii is placed in B. subg. Banksia.

As B. menziesii is not similar to any other Banksia, hybrids are unlikely to occur. The only reported hybrid is a sterile hybrid with 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....

 (Hooker's Banksia), found north of Badgingarra
Badgingarra, Western Australia
Badgingarra is a small town located in the Wheatbelt region of Western Australia, about north of Perth in the Shire of Dandaragan.It lies on the Brand Highway adjacent to the Badgingarra National Park.-History:...

 by Greg Keighery. Manual cross-fertilisation with B. attenuata has resulted in germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

, indicating that these two species are genetically compatible, but natural hybrids are extremely unlikely because the two species flower at different times.

Distribution and habitat

Banksia menziesii grows primarily in deep sandy soils of the Swan Coastal Plain
Swan Coastal Plain
The Swan Coastal Plain in Western Australia is the geographic feature which contains the Swan River as it travels west to the Indian Ocean. The coastal plain continues well beyond the boundaries of the Swan River and its tributaries, as a geological and biological zone, one of Western Australia's...

 and Geraldton Sandplains
Geraldton Sandplains
Geraldton Sandplains is an Interim Biogeographic Regionalisation for Australia region in Western Australia and part of the larger Southwest Australia savanna ecoregion.It has two sub regions: -* Geraldton Hills sub region* Lesuer sub region...

, extending from Waroona
Waroona, Western Australia
Waroona is a town located in the Peel region of Western Australia along the South Western Highway, between Pinjarra and Harvey. The town is the seat of the Shire of Waroona. At the 2006 census, Waroona had a population of 1,864.-History:...

 in the south to Kalbarri
Kalbarri, Western Australia
Kalbarri is a coastal town in the Mid West region located 592 km north of Perth, Western Australia. The town is found at the mouth of the Murchison River and has an elevation of...

 in the north. However, It is uncommon south of Mandurah
Mandurah, Western Australia
Mandurah is the second-largest city in Western Australia and is located approximately south of the state capital, Perth.The city attracts a large number of tourists, including many international visitors...

. It is generally limited to the east by the heavy soils of the Darling Scarp
Darling Scarp
The Darling Scarp, also referred to as the Darling Range or Darling Ranges, is a low escarpment running north-south to the east of the Swan Coastal Plain and Perth, Western Australia...

, but does grow on isolated patches of sand in the Jarrah Forest
Jarrah Forest
Jarrah Forest is an Interim Biogeographic Regionalisation for Australia region in Western Australia.-Location and description:The ecoregion stands on the 300m high Yilgarn block inland plateau and includes wooded valleys such as those of Western Australia's Murray River and the Helena River near...

 and Avon Wheatbelt
Avon Wheatbelt
Avon Wheatbelt is an Interim Biogeographic Regionalisation for Australia region in Western Australia and part of the larger Southwest Australia savanna ecoregion.-Further reading:...

 regions, such as occur near Beverley
Beverley, Western Australia
Beverley is a town located in the Wheatbelt region of Western Australia, south-east of the state capital, Perth, between York and Brookton on the Great Southern Highway...

, Toodyay
Toodyay, Western Australia
Toodyay is a town located in the Wheatbelt region in the Avon Valley, 85 km north-east of Perth, Western Australia. Toodyay is connected to Perth via both rail and road.-History:...

 and Wongan Hills
Wongan Hills, Western Australia
Wongan Hills is a town in the Shire of Wongan-Ballidu, in the Wheatbelt region of Western Australia. The town is approximately 182 km north of the State capital Perth, at an altitude of 286 metres....

. The easternmost known occurrence is a specimen collected by Roger Hnatiuk
Roger Hnatiuk
Roger James Hnatiuk is a Canadian-Australian botanist specialising in biogeography and plant ecology.- Background :Hnatiuk was awarded 1st class honours in botany from the University of Alberta, and went on to graduate with an MSc in plant ecology from the same institution. He was awarded a PhD in...

 in 1979 from north-east of Brookton
Brookton, Western Australia
Brookton is a town located in the Wheatbelt region of Western Australia, Australia, from the state capital, Perth via the Brookton Highway where it crosses the Great Southern Highway. The town is also located on the Great Southern railway line. It is located within, and is the seat of government...

, about 125 km (75 mi) from the coast. Much of its range on the Swan Coastal Plain coincides with Perth's expanding metropolitan area, and much habitat has been lost to clearing.

Together with 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...

 (Candlestick Banksia), B. menziesii is a dominant component in a number of widespread vegetation complexes of the Swan Coastal Plain, including Banksia low woodland and Jarrah-Banksia woodland. These complexes only occur on deep, well-draining sand; in shallower, seasonally wet soils, B. menziesii and B. attenuata give way to other Banksia species such as 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...

 (Swamp Banksia) or 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...

 (Swamp Fox Banksia). On the Geraldton Sandplains to the north, B. menziesii usually occurs as a shrub or small tree emergent above low heath
Heath (habitat)
A heath or heathland is a dwarf-shrub habitat found on mainly low quality acidic soils, characterised by open, low growing woody vegetation, often dominated by plants of the Ericaceae. There are some clear differences between heath and moorland...

.

Ecology

Like many members of the family Proteaceae, Banksia menziesii is largely self-incompatible
Self-incompatibility in plants
Self-incompatibility is a general name for several genetic mechanisms in angiosperms, which prevent self-fertilization and thus encourage outcrossing...

; that is, inflorescences require pollinators to be fertilised and produce seed. One mechanism by which the species promotes cross-pollination with other plants
Allogamy
Allogamy is a term used in the field of biological reproduction describing the fertilization of an ovum from one individual with the spermatozoa of another. By contrast, autogamy is the term used for self-fertilization. In humans, the fertilization event is an instance of allogamy...

 is protandry
Dichogamy
Sequential hermaphroditism is a type of hermaphroditism that occurs in many fish, gastropods and plants. Here, the individual is born one sex and changes sex at some point in their life. They can change from a male to female , or from female to male...

, whereby the male parts release pollen which becomes non-viable before the female parts become receptive on the same flower spike. The individual flowers are uniform, and it is unclear why so few go on to develop follicles. Published in 1988, a field study conducted in banksia woodland near Perth noted that anthesis occurred on an inflorescence at an average rate of 40 to 60 florets opening per day, although this varied widely between different flowerheads. Foraging by honeyeaters would cause the florets to open, but bees would not.

Banksia menziesii provides an important food source, as flowers and seeds, for the threatened
Threatened species
Threatened species are any speciesg animals, plants, fungi, etc.) which are vulnerable to endangerment in the near future.The World Conservation Union is the foremost authority on threatened species, and treats threatened species not as a single category, but as a group of three categories,...

 Short-billed Black Cockatoo (Calyptorhynchus latirostris). Other bird species that have been observed feeding on B. menziesii include the Red-capped Parrot
Red-capped Parrot
The Red-capped Parrot , also called the Pileated Parakeet , and King Parrot locally in Western Australia, is an Australian species of broad-tailed parrot related to the Rosellas.First described by German naturalist Heinrich Kuhl in 1820, from a collection in Albany,...

 (Purpureicephalus spurius), Western Rosella
Western Rosella
The Western Rosella Platycercus icterotis, less commonly known as the Stanley Rosella, Earl of Derby's parakeet or Yellow-cheeked parakeet, is the smallest species of rosella and is found in the South West of Australia. in Eucalypt forests and timbered areas. These are smallish parrots measuring ...

 (Platycercus icterotis), Red-tailed Black Cockatoo
Red-tailed Black Cockatoo
The Red-tailed Black Cockatoo , also known as Banksian- or Banks' Black Cockatoo, is a large cockatoo native to Australia. This species was known as Calyptorhynchus magnificus for many decades until the current scientific name was officially conserved in 1994. It is more common in the drier parts...

 (Calyptorhynchus banksii), Australian Ringneck
Australian Ringneck
The Australian Ringneck is a parrot native to Australia. Except for extreme tropical and highland areas, the species has adapted to all conditions...

 (Barnardius zonarius), Western Gerygone
Western Gerygone
The Western Gerygone is a species of bird in the Acanthizidae family.It is endemic to Australia.Its natural habitats are temperate forests and subtropical or tropical moist lowland forests.-References:...

 (Gerygone fusca) and several honeyeater species, the New Holland Honeyeater
New Holland Honeyeater
The New Holland Honeyeater is a honeyeater species found throughout southern Australia. It was among the first birds to be scientifically described in Australia, and was initially named Certhia novaehollandiae...

 (Phylidonyris novaehollandiae), White-cheeked Honeyeater
White-cheeked Honeyeater
The White-cheeked Honeyeater inhabits the east coast and the south-west corner of Australia. It has a large white patch on its cheek, a brown eye, and a yellow panel on its wing.- Description :...

 (P. nigra), Brown Honeyeater
Brown Honeyeater
The Brown Honeyeater is a honeyeater, a group of birds found mainly in Australia and New Guinea which have highly developed brush-tipped tongues adapted for nectar feeding...

 (Lichmera indistincta), Singing Honeyeater
Singing Honeyeater
The Singing Honeyeater, Lichenostomus virescens is a small bird found in Australia, and is part of the honeyeater family. Although it is common there, it is not very well known in other places....

 (Lichenostomus virescens), Western Spinebill
Western Spinebill
The Western Spinebill, Acanthorhynchus superciliosus, is a honeyeater found in the heath and woodland of south-western Western Australia. It ranges between long, and weighs around . It has a black head, gray back and wings, with a red band behind its neck and from its throat to its breast. There...

 (Acanthorhynchus superciliosus), Red Wattlebird
Red Wattlebird
The Red Wattlebird , also known as Barkingbird or Gillbird, is a honeyeater; a group of birds found mainly in Australia and New Guinea which have highly developed brush-tipped tongues adapted for nectar feeding...

 (Anthochaera carunculata) and Western Wattlebird
Western Wattlebird
The Western Wattlebird is a honeyeater, a passerine bird in the family Meliphagidae. It was formerly lumped with the Little Wattlebird which it closely resembles. It is restricted to south-western Western Australia....

 (A. lunulata). Insects recorded include ants and bees, as well as rove beetle
Rove beetle
The rove beetles are a large family of beetles, primarily distinguished by their short elytra that leave more than half of their abdomens exposed. With over 46,000 species in thousands of genera, the group is the second largest family of beetles after the Curculionidae...

s (family Staphylinidae). A field study south of Perth noted that Banksia menziesii appeared particularly popular with the Brown Honeyeater and Western Spinebill, compared with other banksias.

Twenty-one species from several orders
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of slime molds (myxomycetes) have been isolated from the bark of Banksia menziesii. Over half (13) were from the order Stemonitales, and Echinosteliales and Liceales
Liceales
Liceida is an order of Amoebozoa....

 were also common. The abundance of the first two orders may be due to the acidity of the bark. Another order, the Physarales, was unusually rare—other studies have demonstrated that the order is typically abundant on the bark of various species of tree around the world.

Banksia menziesii regenerates after bushfire by resprouting from its woody lignotuber, or from epicormic buds
Epicormic shoot
An epicormic shoot is a shoot growing from an epicormic bud which lies underneath the bark of a trunk, stem, or branch of a plant.-Epicormic buds:...

 on the trunk. It is generally only weakly serotinous
Serotiny
Serotiny is an ecological adaptation exhibited by some seed plants, in which seed release occurs in response to an environmental trigger, rather than spontaneously at seed maturation. The most common and best studied trigger is fire, and the term serotiny is often used to refer to this specific case...

 in the southern part of its range, that is, it lacks a canopy seed bank
Canopy seed bank
A canopy seed bank or aerial seed bank is the aggregate of viable seed stored by a plant in its canopy. Canopy seed banks occur in plants that postpone seed release for some reason....

 as follicles on old flower spikes in the canopy release their seed after two years, but populations retain more seed as populations move north. Lower canopies and drier climates predispose to hotter fires which are more likely to kill plants and effect seed release, and thus facilitate seedling recruitment.

All banksias have developed proteoid or cluster roots in response to the nutrient poor conditions of Australian soils (particularly lacking in phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

). The plant develop masses of fine lateral roots which form a mat-like structure underneath the soil surface, and enable it to extract nutrients as efficiently possible out of the soil. A study of three co-occurring species in banksia woodland in southwestern Australia—Banksia menziesii, B. attenuata and B. ilicifolia—found that all three develop fresh roots in September after winter rainfall, and that the bacteria populations associated with the root systems of B. menziesii differ from the other two, and that they also change depending on the age of the roots.

Along with Banksia 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...

, Banksia menziesii is a facultative phreatophyte. The two species are less strictly tied to the water table
Water table
The water table is the level at which the submarine pressure is far from atmospheric pressure. It may be conveniently visualized as the 'surface' of the subsurface materials that are saturated with groundwater in a given vicinity. However, saturated conditions may extend above the water table as...

 and hence able to grow in a wider variety of places within banksia woodland habitat around Perth than the co-occurring Banksia 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...

 and Banksia 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...

. Recent falls of the water table on the Swan Coastal Plain from use of the Gnangara Mound
Gnangara Mound
The Gnangara Mound is an area north of Perth, Western Australia where a large mound of sandy soil reaches an elevation of about 60 metres. It stores about 20 cubic kilometres of fresh water, about one hundred times Perth's current annual water usage. It is currently the single most important source...

 aquifer for Perth's water supply as well as years of below average rainfall have caused a drop in the population and vigour of Banksia menziesii since the mid-1960s. A 2009 Spanish study showed Banksia menziesii seedlings to be moderately sensitive to salinity. It is also sensitive to sulfur dioxide
Sulfur dioxide
Sulfur dioxide is the chemical compound with the formula . It is released by volcanoes and in various industrial processes. Since coal and petroleum often contain sulfur compounds, their combustion generates sulfur dioxide unless the sulfur compounds are removed before burning the fuel...

. A 1994 study by Byron Lamont
Byron Lamont
Professor Byron Barnard Lamont is a Western Australian botanist. He is currently a senior researcher within the Department of Environmental Biology of Curtin University of Technology...

 and colleagues from Curtin University
Curtin University of Technology
Curtin University is an Australian university based in Perth, Western Australia, with additional campuses in regional Western Australia and at Miri , Sydney and Singapore...

 found that Banksia menziesii plants within 50 metres (150 ft) of road verges had crowns two and a half times bigger, and set three times as many seeds as plants further away from the road, and that this was likely due to increased availability of nutrients and water from runoff
Surface runoff
Surface runoff is the water flow that occurs when soil is infiltrated to full capacity and excess water from rain, meltwater, or other sources flows over the land. This is a major component of the water cycle. Runoff that occurs on surfaces before reaching a channel is also called a nonpoint source...

.

Cultivation

Banksia menziesii has several horticultural features including patterned buds and follicles, attractive pinkish new growth, and showy flowerheads. A dwarf form is commonly sold in nurseries. Byron Lamont has observed that dwarf plants may grow into taller single-trunked plants in cultivation.

The plant is fairly easy to grow in a mediterranean climate
Mediterranean climate
A Mediterranean climate is the climate typical of most of the lands in the Mediterranean Basin, and is a particular variety of subtropical climate...

 with good drainage and a light (sandy) soil; however, with medium to high susceptibility to Phytophthora cinnamomi
Phytophthora cinnamomi
Phytophthora cinnamomi is a soil-borne water mould that produces an infection which causes a condition in plants called root rot or dieback. The plant pathogen is one of the world's most invasive species and is present in over 70 countries from around the world.- Life cycle and effects on plants :P...

 dieback, it is unreliable in conditions which favour the growth of the soil-borne water mould, such as summer humidity or poor drainage. The use of phosphite
Phosphorous acid
Phosphorous acid is the compound described by the formula H3PO3. This acid is diprotic , not triprotic as might be suggested by this formula. Phosphorous acid is as an intermediate in the preparation of other phosphorus compounds.-Nomenclature and tautomerism:H3PO3 is more clearly described with...

 fungicides does reduce the spread of Phytophthora. It generally does poorly in eastern Australia, although it is grown commercially in southeastern South Australia as a cut flower
Floriculture
Floriculture, or flower farming, is a discipline of horticulture concerned with the cultivation of flowering and ornamental plants for gardens and for floristry, comprising the floral industry...

 crop. It is also grown in California and Hawaii.

Seeds do not require any treatment prior to planting, and take 26 to 40 days to germinate. Seeds that grow into yellow-flowered plants are pale and unpigmented, while future bronze- and red-flowered plants are dark greyish and black respectively. It generally takes about five to seven years to flower from seed. The plant favours sandy well-drained soils and a sunny position and can be heavily pruned
Pruning
Pruning is a horticultural practice involving the selective removal of parts of a plant, such as branches, buds, or roots. Reasons to prune plants include deadwood removal, shaping , improving or maintaining health, reducing risk from falling branches, preparing nursery specimens for...

 if necessary, as new growth can arise from the lignotuber. Although it is readily propagated by seed, experiments with in vitro propagation found Banksia menziesii to be more difficult than other species trialled.

Art and culture

The nectar of Banksia menziesii was used in a drink by the Beeloo Whadjug Nyoongar, who were also called the Mungyt people. Along with beverages from other species, it was drunk at special sweet water festivals. Banksia menziesii was the subject of a book by botanical artist Philippa Nikulinsky
Philippa Nikulinsky
Philippa Nikulinsky, born in 1942, is an artist and botanical illustrator based in Western Australia.-Biography:Nikulinsky was born in Kalgoorlie in 1942, a remote region in central Western Australia. She began working as an illustrator of natural history in the mid 1970s, specialising in plants...

, which showed the progress of an inflorescence from bud through flowering to fruiting and seed release over 22 watercolour plates. Noted wildflower artist Ellis Rowan
Ellis Rowan
Marian Ellis Rowan , known as Ellis Rowan, was a well-known Australian botanical illustrator. She also did series of illustrations on birds, butterflies and insects....

 also painted it. It was one of several wildflowers depicted on a series of plates produced by the British pottery firm Wedgwood
Wedgwood
Wedgwood, strictly speaking Josiah Wedgwood and Sons, is a pottery firm owned by KPS Capital Partners, a private equity company based in New York City, USA. Wedgwood was founded on May 1, 1759 by Josiah Wedgwood and in 1987 merged with Waterford Crystal to create Waterford Wedgwood, an...

in the early 1990s.

External links

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