Dryandra Sessilis
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''Banksia sessilis'', commonly known as parrot bush, is a species of
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
or tree in the plant genus '' Banksia'' of the family
Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...
. It had been known as ''Dryandra sessilis'' until 2007, when the genus ''
Dryandra ''Banksia'' ser. ''Dryandra'' is a series of 94 species of shrub to small tree in the plant genus ''Banksia''. It was considered a separate genus named ''Dryandra'' until early 2007, when it was merged into ''Banksia'' on the basis of extensiv ...
'' was sunk into ''Banksia''. The Noongar peoples know the plant as budjan or butyak. Widespread throughout southwest Western Australia, it is found on sandy soils over
laterite Laterite is both a soil and a rock type rich in iron and aluminium and is commonly considered to have formed in hot and wet tropical areas. Nearly all laterites are of rusty-red coloration, because of high iron oxide content. They develop by ...
or limestone, often as an understorey plant in open forest, woodland or shrubland. Encountered as a shrub or small tree up to in height, it has prickly dark green leaves and dome-shaped cream-yellow
flowerheads A pseudanthium (Greek for "false flower"; ) is an inflorescence that resembles a flower. The word is sometimes used for other structures that are neither a true flower nor a true inflorescence. Examples of pseudanthia include flower heads, compos ...
. Flowering from winter through to late spring, it provides a key source of food—both the nectar and the insects it attracts—for
honeyeater The honeyeaters are a large and diverse family (biology), family, Meliphagidae, of small to medium-sized birds. The family includes the Epthianura, Australian chats, myzomelas, friarbirds, wattlebirds, Manorina, miners and melidectes. They are ...
s in the cooler months, and species diversity is reduced in areas where there is little or no parrot bush occurring. Several species of honeyeater, some species of native bee, and the European honey bee seek out and consume the nectar, while the
long-billed black cockatoo Baudin's black cockatoo (''Zanda baudinii''), also known as Baudin's cockatoo or the long-billed black cockatoo, is a species of genus '' Zanda'' found in southwest Australia. The epithet commemorates the French explorer Nicolas Baudin. It has ...
and Australian ringneck eat the seed. The life cycle of ''Banksia sessilis'' is adapted to regular bushfires. Killed by fire and regenerating by seed afterwards, each shrub generally produces many flowerheads and a massive amount of seed. It can recolonise disturbed areas, and may grow in thickets. ''Banksia sessilis'' has a somewhat complicated
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
history. It was collected from King George Sound in 1801 and described by Robert Brown in 1810 as ''Dryandra floribunda'', a name by which it was known for many years. However, Joseph Knight had published the name ''Josephia sessilis'' in 1809, which had precedence due to its earlier date, and the specific name was formalised in 1924. Four varieties are recognised. It is a prickly plant with little apparent horticultural potential; none of the varieties are commonly seen in cultivation. A profuse producer of nectar, ''B. sessilis'' is valuable to the
beekeeping Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus '' Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees are also kept. ...
industry.


Description

''Banksia sessilis'' grows as an upright shrub or small tree up to high, without a lignotuber. In most varieties, new stems are covered in soft, fine hairs that are lost with maturity; but new stems of ''B. sessilis'' var. ''flabellifolia'' are usually hairless. Leaves are blue-green or dark green. Their shape differs by variety: in var. ''cygnorum'' and var. ''flabellifolia'' they are wedge-shaped, with teeth only near the apex; in var. ''cordata'' they are wedge-shaped, but with teeth along the entire margin; and in var. ''sessilis'' they are somewhat broader at the base, sometimes almost oblong in shape. Leaf size ranges from in length, and in width. They may be sessile (that is, growing directly from the stem without a petiole) or on a petiole up to long. The inflorescences are cream or yellow, and occur in domed
heads A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may no ...
wide, situated at the end of a stem. Each head contains from 55 to 125 individual flowers, surrounded at the base by a whorl of short
involucral bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
s. As with most other
Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...
, individual flowers consist of a tubular perianth made up of four united tepals, and one long wiry style. The style end is initially trapped inside the upper perianth parts, but breaks free at anthesis. In ''B. sessilis'' the perianth is straight, long, and pale yellow. The style is slightly shorter, also straight, and cream-coloured. Thus in ''B. sessilis'', unlike many other ''Banksia'' species, the release of the style at anthesis does not result in a showy flower colour change. One field study found that anthesis took place over four days, with the outer flowers opening first and moving inwards. Flowering mostly takes place from July to November; var. ''sessilis'' can start as early as May. After flowering, the flower parts wither and fall away, and up to four follicles develop in the receptacle (the base of the flower head). Young follicles are covered in a fine fur, but this is lost as they mature. Mature follicles are
ovoid An oval () is a closed curve in a plane which resembles the outline of an egg. The term is not very specific, but in some areas (projective geometry, technical drawing, etc.) it is given a more precise definition, which may include either one or ...
in shape, and measure in length. Most follicles open as soon as they are ripe, revealing their contents: a woody seed separator and up to two winged seeds.


Discovery and naming

Specimens of ''B. sessilis'' were first collected by Scottish surgeon Archibald Menzies during the visit of the Vancouver Expedition to King George Sound in September and October 1791. No firm location or collection date can be ascribed to Menzies' specimens, as their labels simply read "New Holland, King Georges Sound, Mr. Arch. Menzies", and Menzies' journal indicates that he collected over a wide area, visiting a different location every day from 29 September to 8October. In addition to ''B. sessilis'', Menzies collected plant material of '' B. pellaeifolia'', and seeds of at least four more ''Banksia'' species. This was therefore an important early collection for the genus, only seven species of which had previously been collected. Menzies' seed specimens were sent to England from
Sydney Sydney ( ) is the capital city of the state of New South Wales, and the most populous city in both Australia and Oceania. Located on Australia's east coast, the metropolis surrounds Sydney Harbour and extends about towards the Blue Mountain ...
in 1793, but his plant material remained with him for the duration of the voyage, during which some material was lost. On his return to England in 1795, the surviving specimens were deposited into the
herbarium A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...
of Sir Joseph Banks, where they lay undescribed for many years. The next collection was made in December 1801, when King George Sound was visited by HMS ''Investigator'' under the command of
Matthew Flinders Captain Matthew Flinders (16 March 1774 – 19 July 1814) was a British navigator and cartographer who led the first inshore circumnavigation of mainland Australia, then called New Holland. He is also credited as being the first person to u ...
. On board were
botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
Robert Brown, botanical artist Ferdinand Bauer, and gardener Peter Good. All three men gathered material for Brown's specimen collection, including specimens of ''B. sessilis'', but neither Brown's nor Good's diary can be used to assign a precise location or date for their discovery of the species. Good also made a separate seed collection, which included ''B. sessilis'', and the species was drawn by Bauer. Like nearly all of his field drawings of Proteaceae, Bauer's original field sketch of ''B. sessilis'' was destroyed in a
Hofburg fire The term "Hofburg fire" "Pervasive 2004, April 21–23, Linz / Vienna, Austria" (Hofburg history), Pervasive 2004, April 2004, Pervasive2004.org webpage: pervasive2004-Hofburg-fire refers to any of several major fires that burned in ...
in 1945. A painting based on the drawing survives, however, at the Natural History Museum in London. On returning to England in 1805, Brown began preparing an account of his Australian plant specimens. In September 1808, with Brown's account still far from finished, Swedish botanist
Jonas Dryander Jonas Carlsson Dryander (5 March 1748 – 19 October 1810) was a Swedish botanist. Biography Dryander was born in Gothenburg, Sweden. He was the son of Carl Leonard Dryander and Brita Maria Montin. He was a pupil of Carl Linnaeus at Uppsala ...
asked him to prepare a separate paper on the
Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...
so he could use the genera erected by Brown in a new edition of '' Hortus Kewensis''. Brown immediately began a study of the Proteaceae, and in January 1809 he read to the Linnean Society of London a
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
on the family entitled '' On the Proteaceae of Jussieu''. Among the eighteen new genera presented was one that Brown named ''Josephia'' in honour of Banks. Brown's paper was approved for printing in May 1809, but did not appear in print until March the following year. In the meantime, Joseph Knight published '' On the cultivation of the plants belonging to the natural order of Proteeae'', which appeared to draw heavily on Brown's unpublished material, without permission, and in most cases without attribution. It contained the first publication of Brown's ''Josephia'', for which two species were listed. The first, ''Josephia sessilis'', was based on one of Menzies' specimens: "This species, discovered by Mr. A. Menzies on the West coast of New Holland, is not unlike some varieties of ''
Ilex aquifolium ''Ilex aquifolium'', the holly, common holly, English holly, European holly, or occasionally Christmas holly, is a species of flowering plant in the family (botany), family Aquifoliaceae, native plant, native to western and southern Europe, nort ...
'', and now in his Majesty's collection at Kew." The etymology of the specific epithet was not explicitly stated, but it is universally accepted that it comes from the Latin ''sessilis'' ( sessile, stalkless), in reference to the sessile leaves of this species. Blame for the alleged
plagiarism Plagiarism is the fraudulent representation of another person's language, thoughts, ideas, or expressions as one's own original work.From the 1995 '' Random House Compact Unabridged Dictionary'': use or close imitation of the language and thought ...
largely fell on Richard Salisbury, who had been present at Brown's readings and is thought to have provided much of the material for Knight's book. Salisbury was ostracized by the botanical community, which undertook to ignore his work as much as possible. By the time Brown's monograph appeared in print, Brown had exchanged the generic name ''Josephia'' for ''Dryandra'', giving the name ''Dryandra floribunda'' to Knight's ''Josephia sessilis''. As there were then no firm rules pertaining to priority of publication, Brown's name was accepted, and remained the current name for over a century. Another significant early collection was the apparent discovery of the species at the Swan River in 1827. In that year, the colonial botanist of New South Wales Charles Fraser visited the area as part of an exploring expedition under James Stirling. Among the plants Fraser found growing on the south side of the river entrance was "a beautiful species of Dryandra", which was probably this species. Over the course of the 19th century, the principle of priority in naming gradually came to be accepted by botanists, as did the need for a mechanism by which names in current usage could be conserved against archaic or obscure prior names. By the 1920s, ''Dryandra'' R.Br. was effectively conserved against ''Josephia'' Knight; a mechanism for formal conservation was put in place in 1933. Brown's specific name, however, was not conserved, and Karel Domin overturned ''Dryandra floribunda'' R.Br. by transferring Knight's name into ''Dryandra'' as ''Dryandra sessilis'' (Knight) Domin in 1924. This name was current until 2007, when all ''Dryandra'' species were transferred into '' Banksia'' by Austin Mast and Kevin Thiele. The full citation for the current name is thus ''Banksia sessilis'' (Knight) A.R.Mast & K.R.Thiele.


Common names

The first common names for this species were literal translations of the scientific names. When published as ''Josephia sessilis'' in 1809, it was given the common name ''sessile Josephia''. Brown did not offer a common name when he published ''Dryandra floribunda'' in 1810, but later that year the '' Hortus Kewensis'' translated it as ''many flowered dryandra''. This name was also used when the plant was featured in '' Curtis's Botanical Magazine'' in 1813. In Australia, the names ''prickly banksia'' and ''shaving-brush flower'' were offered up by Emily Pelloe in 1921, the latter because "when in bud the flower very much resembles a shaving-brush". ''Shaving-brush flower'' was still in use as late as the 1950s. The name ''holly-leaved dryandra'' was used when the plant was featured as part of a series of articles in the '' Western Mail'' of 1933–34, and this was taken up by William Blackall in 1954, and was still in use as late as 1970. Meanwhile, Gardner used the name ''parrot bush'' in 1959, a name derived from the observation that the blooms attract parrots, by which the species was already "well-known to bee-keepers". This name was widely adopted, and since 1970 has been in almost exclusive usage. The only indigenous names reported for the plant are ''Budjan'' and ''But-yak''. These were published by Ian Abbott in his 1983 ''Aboriginal Names for Plant Species in South-western Australia'', with Abbott suggesting that the latter name should be preferred, but with the orthography "Pudjak". However, Abbott sources these names to George Fletcher Moore's 1842 '' A Descriptive Vocabulary of the Language of the Aborigines'', which in fact attributes these names to the species ''Dryandra fraseri'' (now ''
Banksia fraseri ''Banksia fraseri'' is a species of shrub that is endemic to Western Australia. It has hairy stems, broadly linear pinnatisect leaves with between four and eighteen sharply-pointed lobes on each side, between eighty and one hundred pink to cre ...
''). It is unclear whether Abbott has corrected Moore's error, or introduced an error of his own.


Taxonomy


Infrageneric placement

Brown's 1810 monograph did not include an infrageneric classification of ''Dryandra'', and neither did his '' Prodromus'', published later that year. In 1830, however, he introduced the first taxonomic arrangement of ''Dryandra'', placing ''D. floribunda'' in section ''
Dryandra verae ''Dryandra'' subg. ''Dryandra'' is an obsolete clade of plant. It was a series within the former genus ''Dryandra'' (now ''Banksia'' ser. ''Dryandra''). The name was first published at sectional rank as ''Dryandra verae'' in 1830, before bein ...
'' along with most other species, because its follicles contain a single seed separator. ''Dryandra verae'' was renamed ''Eudryandra'' by Carl Meissner in 1845. Eleven years later Meissner published a new arrangement, retaining ''D. floribunda'' in ''D.'' sect. ''Eudryandra'', and further placing it in the unranked subgroup § ''Ilicinae'', because of the similarity of its leaves to those of '' Ilex'' (holly). In 1870, George Bentham published a revised arrangement in his '' Flora Australiensis''. Bentham retained section ''Eudryandra'', but abandoned almost all of Meissner's unranked groups, including § ''Ilicinae''. ''D. floribunda'' was instead placed in ''D.'' ser. ''Floribundae'' along with four other species with small, mostly terminal flowers, left exposed by their having unusually short floral leaves. Bentham's arrangement stood for over a hundred years, eventually replaced in 1996 by the arrangement of
Alex George Alexander or Alex George may refer to: *Alex George (botanist) (born 1939), Australian botanist * Alexander L. George (1920–2006), American political scientist * Alexander George (philosopher), American philosopher *Alex George (motorcyclist), Sc ...
. Section ''Eudryandra'' was promoted to subgenus rank, but replaced by the autonym ''D.'' subg. ''Dryandra''. ''D. sessilis'', as this species was now called, was retained in ''D.'' ser. ''Floribundae'', but alone, as the series was redefined as containing only those taxa that apparently lack floral bracts altogether. The placement of ''D. sessilis'' in George's arrangement, with 1999 and 2005 amendments, may be summarised as follows: :''Dryandra'' (now ''Banksia'' ser. ''Dryandra'') :: ''D.'' subg. ''Dryandra'' ::: ''D.'' ser. ''Floribundae'' ::::''D. sessilis'' (now ''B. sessilis'') :::::''D. sessilis'' var. ''sessilis'' (now ''B. sessilis'' var. ''sessilis'') :::::''D. sessilis'' var. ''flabellifolia'' (now ''B. sessilis'' var. ''flabellifolia'') :::::''D. sessilis'' var. ''cordata'' (now ''B. sessilis'' var. ''cordata'') :::::''D. sessilis'' var. ''cygnorum'' (now ''B. sessilis'' var. ''cygnorum'') ::: ''D.'' ser. ''Armatae'' ::: ''D.'' ser. ''Marginatae'' ::: ''D.'' ser. ''Folliculosae'' ::: ''D.'' ser. ''Acrodontae'' ::: ''D.'' ser. ''Capitellatae'' ::: ''D.'' ser. ''Ilicinae'' ::: ''D.'' ser. ''Dryandra'' ::: ''D.'' ser. ''Foliosae'' ::: ''D.'' ser. ''Decurrentes'' ::: ''D.'' ser. ''Tenuifoliae'' ::: ''D.'' ser. ''Runcinatae'' ::: ''D.'' ser. ''Triangulares'' ::: ''D.'' ser. ''Aphragma'' ::: ''D.'' ser. ''Ionthocarpae'' ::: ''D.'' ser. ''Inusitatae'' ::: ''D.'' ser. ''Subulatae'' ::: ''D.'' ser. ''Gymnocephalae'' ::: ''D.'' ser. ''Plumosae'' ::: ''D.'' ser. ''Concinnae'' ::: ''D.'' ser. ''Obvallatae'' ::: ''D.'' ser. ''Pectinatae'' ::: ''D.'' ser. ''Acuminatae'' ::: ''D.'' ser. ''Niveae'' :: ''D.'' subg. ''Hemiclidia'' :: ''D.'' subg. ''Diplophragma'' George's arrangement remained current until 2007, when Austin Mast and Kevin Thiele transferred ''Dryandra'' into ''Banksia''. They also published ''B.'' subg. ''Spathulatae'' for the ''Banksia'' taxa having spoon-shaped cotyledons, thus redefining ''B.'' subg. ''Banksia'' as comprising those that do not. They were not ready, however, to tender an infrageneric arrangement encompassing ''Dryandra'', so as an interim measure they transferred ''Dryandra'' into ''Banksia'' at series rank. This minimised the nomenclatural disruption of the transfer, but also caused George's rich infrageneric arrangement to be set aside. Thus under the interim arrangements implemented by Mast and Thiele, ''B. sessilis'' is placed in ''B.'' subg. ''Banksia'', ser. ''Dryandra''.


Varieties

Four varieties are recognised: * ''B. sessilis'' var. ''sessilis'' is an autonym that encompasses the type material of the species. This is the most widespread variety, occurring from Regans Ford and Moora in the north, south-east to Albany, and inland as far as Wongan Hills, Pingelly and
Kulin Kulin may refer to: Places *Kulin, Western Australia, a small town in Australia ** Shire of Kulin, a local government area *Kulin, Iran, a village near Tehran *Kulin, Lower Silesian Voivodeship, a village in south-west Poland *Kulin, Kuyavian-Pome ...
. Its blue-green leaves are cuneate (wedge-shaped) or oblong, and are usually two to three centimetres long but may reach five. * ''B. sessilis'' var. ''cordata'' was published as ''Dryandra floribunda'' var. ''cordata'' by Carl Meissner in 1848. In 1870, George Bentham published ''D. floribunda'' var. ''major'', but this is now considered a taxonomic synonym of ''B. sessilis'' var. ''cordata''. It has larger inflorescences than var. ''sessilis'', as well as larger dark green, rather than blue green leaves. It is found in the state's far southwest, between Capes Leeuwin and Naturaliste, and east to Walpole, and grows on sandy soils over limestone. * ''B. sessilis'' var. ''cygnorum'' has its roots in Michel Gandoger's publication of two new species names in 1919. He published ''Dryandra cygnorum'' and ''Dryandra quinquedentata'', but in 1996 both of these were found to refer to the same taxon, which
Alex George Alexander or Alex George may refer to: *Alex George (botanist) (born 1939), Australian botanist * Alexander L. George (1920–2006), American political scientist * Alexander George (philosopher), American philosopher *Alex George (motorcyclist), Sc ...
gave variety rank as ''Dryandra sessilis'' var. ''cygnorum''. The term ''cygnorum'' is Latin for "swan" and relates to the Swan River, which runs past the suburb of Melville where the type material was collected. It has smaller dark green leaves only long and wide, whose teeth are limited to the distal part of the leaf. The range is along the Western Australian coastline from Dongara southwards past
Fremantle Fremantle () () is a port city in Western Australia, located at the mouth of the Swan River in the metropolitan area of Perth, the state capital. Fremantle Harbour serves as the port of Perth. The Western Australian vernacular diminutive for ...
, and east to Lake Indoon and Kings Park. * ''B. sessilis'' var. ''flabellifolia'' was published by George in 1996, the type specimen having been collected northwest of
Northampton Northampton () is a market town and civil parish in the East Midlands of England, on the River Nene, north-west of London and south-east of Birmingham. The county town of Northamptonshire, Northampton is one of the largest towns in England; ...
in 1993. The northernmost of the four varieties, it is found from Kalbarri south to Geraldton and Northampton. There are some scattered records further south towards Moora. Its specific name is derived from the Latin ''flabellum'' "fan" and ''folium'' "leaf". Its leaves are fan shaped, with a long, toothless lower margin, and a toothed end. Its stems are hairless, unlike the other varieties.


Distribution and habitat

''Banksia sessilis'' is endemic to the Southwest Botanical Province, a floristic province renowned as a biodiversity hotspot, located in the southwest corner of Western Australia. This area has a Mediterranean climate, with wet winters and hot, dry summers. ''B. sessilis'' occurs throughout much of the province, ranging from Kalbarri in the north, south to Cape Leeuwin, east along the south coast as far as Bremer Bay, and inland to Wongan Hills and
Kulin Kulin may refer to: Places *Kulin, Western Australia, a small town in Australia ** Shire of Kulin, a local government area *Kulin, Iran, a village near Tehran *Kulin, Lower Silesian Voivodeship, a village in south-west Poland *Kulin, Kuyavian-Pome ...
. It thus spans a wide range of climates, occurring in all but the semi-arid areas well inland. It is also absent from the Karri forest in the cool, wet, southwest corner of the province, but even there, ''B. sessilis'' var. ''cordata'' occurs along the coast. The species tolerates a range of soils, requiring only that its soil be well-drained. Like most dryandras, it grows well in
lateritic Laterite is both a soil and a rock type rich in iron and aluminium and is commonly considered to have formed in hot and wet tropical areas. Nearly all laterites are of rusty-red coloration, because of high iron oxide content. They develop by ...
soils and gravels; this species is also found in deep sand, sand over laterite, and sand over limestone. It also occurs in a range of vegetation complexes, including coastal and kwongan heath, tall
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It m ...
, woodland and open forest. It is a common understorey plant in drier areas of Jarrah forest, and forms thickets on limestone soils of the
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 geol ...
. ''Banksia sessilis'' sets a large amount of seed and is an aggressive coloniser of disturbed and open areas; for example, it has been recorded colonising gravel pits in the Darling Scarp. Nothing is known of the conditions that affect its distribution, as its
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
is as yet unstudied. An assessment of the potential
impact of climate change The effects of climate change impact the physical environment, ecosystems and human societies. The environmental effects of climate change are broad and far-reaching. They affect the water cycle, oceans, sea and land ice (glaciers), sea level ...
on this species found that its range is likely to contract by half in the face of severe change, but unlikely to change much under less severe scenarios.


Ecology


As food

The
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
of ''B. sessilis'' is an important component of the diet of several species of
honeyeater The honeyeaters are a large and diverse family (biology), family, Meliphagidae, of small to medium-sized birds. The family includes the Epthianura, Australian chats, myzomelas, friarbirds, wattlebirds, Manorina, miners and melidectes. They are ...
. In one study, ''B. sessilis'' was found to be the main source of nectar for all six species studied, namely the
tawny-crowned honeyeater The tawny-crowned honeyeater (''Gliciphila melanops'') is a passerine bird native to southern Australia. Taxonomy The tawny-crowned honeyeater was originally described by ornithologist John Latham in 1801 as ''Certhia melanops''. Its specific e ...
(''Gliciphila melanops''),
white-cheeked honeyeater The white-cheeked honeyeater (''Phylidonyris niger'') inhabits the east coast and the south-west corner of Australia. It has a large white patch on its cheek, brown eyes, and a yellow panel on its wing. Taxonomy The white-cheeked honeyeater was ...
(''Phylidonyris niger''), western spinebill (''Acanthorhynchus superciliosus''), brown honeyeater (''Lichmera indistincta''),
brown-headed honeyeater The brown-headed honeyeater (''Melithreptus brevirostris'') is a species of passerine bird in the family Meliphagidae. It is endemic to Australia. Its natural habitats are temperate forests and Mediterranean-type shrubby vegetation. Taxonomy T ...
(''Melithreptus brevirostris''), and
black honeyeater The black honeyeater (''Sugomel nigrum'') is a species of bird in the honeyeater family Meliphagidae. The black honeyeater exhibits sexual dimorphism, with the male being black and white while the female is a speckled grey-brown; immature birds l ...
(''Certhionyx niger''). Moreover, ''B. sessilis'' played an important role in their distributions, with species that feed only on nectar occurring only where ''B. sessilis'' occurs, and remaining for longest at sites where ''B. sessilis'' is most abundant. Other honeyeaters that have been recorded feeding on ''B. sessilis'' include the red wattlebird (''Anthochaera carunculata''), western wattlebird (''A. lunulata''), and
New Holland honeyeater The New Holland honeyeater (''Phylidonyris novaehollandiae'') 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 novaehollandi ...
(''Phylidonyris novaehollandiae''). Furthermore, a study of bird species diversity in
wandoo Wandoo is the common name for a number of Western Australian ''Eucalyptus'' species, all of which have smooth white bark. The original "wandoo" is ''Eucalyptus wandoo''. Additional species have been given this name because of a perceived likeness w ...
woodland around Bakers Hill found that honeyeater species and numbers were much reduced in forest that lacked a ''Banksia sessilis'' understory; the plant is a key source of nectar and insects during the winter months. A field study in jarrah forest 9 km south of Jarrahdale, where ''B. sessilis'' grows in scattered clumps, found that western wattlebirds and New Holland honeyeaters sought out groups of plants with the greatest numbers of new inflorescences, particularly those one or two days after anthesis, where nectar yield was highest. The birds likely recognises these by visual clues. ''Banksia sessilis'' is also a source of food for the Australian ringneck (''Barnardius zonarius''), and the
long-billed black cockatoo Baudin's black cockatoo (''Zanda baudinii''), also known as Baudin's cockatoo or the long-billed black cockatoo, is a species of genus '' Zanda'' found in southwest Australia. The epithet commemorates the French explorer Nicolas Baudin. It has ...
(''Calyptorhynchus baudinii''), which tear open the follicles and consume the seeds. The introduced European honey bee (''Apis mellifera'') has also been observed feeding on ''B. sessilis'', as have seven species of
native bee Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entertai ...
, comprising four species of '' Hylaeus'' (including the
banksia bee ''Hylaeus alcyoneus'', commonly known as the banksia bee, is a bee species endemic to Australia where it is commonly found in the coastal heaths of eastern and southern Western Australia. This bee is an important pollinator of ''Banksia'' species ...
''H. alcyoneus''), two of ''
Leioproctus ''Leioproctus'' is a genus in the plaster bee family Colletidae. Its members are primarily found in Australasia and temperate South America, and include the most common native bees in New Zealand. Species It includes the following species: *' ...
'', and a '' Lasioglossum''.


Life cycle

Honeyeaters are clearly the most important pollination vector, as inflorescences from which honeyeaters are excluded generally do not set any fruit. Moreover, honeyeaters have been observed moving from tree to tree with significant loads of ''B. sessilis'' pollen on their foreheads, beaks and throats, having acquired it by brushing against pollen presenters while foraging for nectar; experiments have shown that some of this pollen may be subsequently deposited on stigmas during later foraging. The flowers of ''B. sessilis'' have adaptations that encourage outcrossing. Firstly, they are protandrous: a flower's pollen is released around 72 hours before it becomes itself receptive to pollen, by which time around half of its pollen has lost its viability. Secondly, the period of maximum nectar production closely matches the period during which the flower is sexually active, so honeyeaters are enticed to visit at the most opportune time for pollination. This has proven an effective strategy: almost all pollen is removed within two to three hours of presentation. In addition, honeyeaters tend to move between inflorescences on different plants, rather than between inflorescences on the same plant, at least in high density sites. These factors combine to make it fairly unusual for a flower to be fertilised by its own pollen. When self-fertilisation does occur, whether autogamous or
geitonogamous Geitonogamy (from Greek ''geiton'' (γείτων) = neighbor + ''gamein'' (γαμεῖν) = to marry) is a type of self-pollination. Geitonogamous pollination is sometimes distinguished from the fertilizations that can result from it, geitonogamy. ...
, the resulting seed is almost always aborted, and the species ultimately achieves an outcrossing rate of nearly 100%, at least in high density sites. Limited data for low-density sites, where honeyeaters move from plant to plant less frequently, suggest more of a
mixed-mating system The mixed-mating model is a mathematical model that describes the mating system of a plant population in terms of degree of self-fertilisation. It is a fairly simplistic model, employing several simplifying assumptions, most notably the assumption t ...
. The species is a prolific flowerer, and this, combined with the very high outcrossing rates, results in massive seed output. In one study, the average number of seeds produced per ''B. sessilis'' plant was 622, compared with an average of two for '' B. dallanneyi''. This exceptionally high fecundity can be understood as an adaption to regular bushfire. Most ''Banksia'' species can be placed in one of two broad groups according to their response to fire: '' resprouters'' survive fire, resprouting from a lignotuber or, more rarely,
epicormic buds 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 lie dormant beneath the bark, their growth suppressed by hormones from active shoots higher up ...
protected by thick bark; ''reseeders'' are killed by fire, but populations are rapidly re-established through the recruitment of seedlings. ''B. sessilis'' is a reseeder, but it differs from many other reseeders in not being strongly serotinous: the vast majority of seeds are released spontaneously in autumn, even in the absence of fire. The degree of serotiny is a matter of some contradiction in the scientific literature: it has been treated as "serotinous", "weakly serotinous" and "non-serotinous". Regardless of the terminology used, the massive spontaneous seed output of ''B. sessilis'' is its primary survival strategy, and is so effective the species has a reputation as an excellent coloniser. However, this strategy, together with its relatively long juvenile period, makes it vulnerable to overly frequent fire. Seeds of ''B. sessilis'' are short-lived, and must germinate in the winter following their release, or they die. They are also very sensitive to heating, and thus killed by bushfire; in one study, just 30 seconds in boiling water reduced the germination rate from 85% to 22%, and not a single seed survived one minute of boiling. Like most other
Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...
, ''B. sessilis'' has compound cluster roots, roots with dense clusters of short lateral rootlets that form a mat in the soil just below the leaf litter. These exude a range of carboxylates, including citrate, malonate and ''trans''-aconitate, that act as
acid phosphatase Acid phosphatase (EC 3.1.3.2, acid phosphomonoesterase', phosphomonoesterase, glycerophosphatase, acid monophosphatase, acid phosphohydrolase, acid phosphomonoester hydrolase, uteroferrin, acid nucleoside diphosphate phosphatase, orthophosphoric-m ...
, allowing the absorption of nutrients from nutrient-poor soils, such as the phosphorus-deficient native soils of
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
.


Disease

''Banksia sessilis'' is highly susceptible to dieback caused by the introduced plant pathogen '' Phytophthora cinnamomi'', a soil-borne water mould that causes
root rot Root rot is a condition in which anoxic conditions in the soil or potting media around the roots of a plant cause them to rot. This occurs due to excessive standing water around the roots. It is found in both indoor and outdoor plants, although ...
; in fact it is so reliably susceptible it is considered a good
indicator species A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
for the presence of the disease. Most highly susceptible species quickly become locally extinct in infected areas, and in the absence of hosts the disease itself eventually dies out. However, ''B. sessilis'', being an aggressive coloniser of disturbed and open ground, often colonises old disease sites. The new colonies are themselves infected, and thus ''P. cinnamomi'' survives at these sites indefinitely. The application of phosphite inhibits growth of ''P. cinnamomi'' in ''B. sessilis'', but does not kill the pathogen. In one study, a foliar spray containing phosphite inhibited the growth of ''P. cinnamomi'' by over 90% in plants infected with ''B. sessilis'' two weeks after spraying, and by 66% in plants infected one year after spraying; yet most plants infected shortly before or after spraying were dead 100 days later, while nearly all plants infected seven months later spraying survived a further 100 days. Phosphite is not known to affect plant growth, but has been shown to reduce pollen fertility: one study recorded fertility reductions of up to 50%, and, in a separate experiment, fertility reductions that persisted for more than a year. Infection of coastal stands of ''B. sessilis'' by the fungus ''
Armillaria luteobubalina ''Armillaria luteobubalina'', commonly known as the Australian honey fungus, is a species of mushroom in the family Physalacriaceae. Widely distributed in southern Australia, the fungus is responsible for a disease known as ''Armillaria'' root ...
'' has also been recorded. The
apparent infection rate Apparent infection rate is an estimate of the rate of progress of a disease, based on proportional measures of the extent of infection at different times. Firstly, a proportional measure of the extent of infection is chosen as the disease extent ...
of 0.31 is quite slow compared to the progress of other '' Armillaria'' species through pine plantations.


Cultivation


History

It is not known whether the seed collection sent to the Royal Botanic Gardens, Kew, by Menzies in 1793 included seeds of ''B. sessilis'', but if it did then it did not germinate. The species was successfully germinated, however, from Good's seed, which was sent from Sydney on 6June 1802 and arrived at Kew the following year. According to Brown's notes it was flowering at Kew by May 1806, and in 1810 it was reported in the second edition of ''Hortus Kewensis'' as flowering "most part of the Year". In 1813 a flowering specimen from the nursery of Malcolm and Sweet was featured as Plate 1581 in '' Curtis's Botanical Magazine''. By the 1830s the species was in cultivation in continental Europe. It was recorded as being cultivated in the garden of
Karl von Hügel Karl may refer to: People * Karl (given name), including a list of people and characters with the name * Karl der Große, commonly known in English as Charlemagne * Karl Marx, German philosopher and political writer * Karl of Austria, last Austrian ...
in Vienna, Austria in 1831, and in 1833 it was listed amongst the rare plants that had been introduced into Belgium. Along with several hundred other native Australian plants it was exhibited at plant shows held at Utrecht and
Haarlem Haarlem (; predecessor of ''Harlem'' in English) is a city and municipality in the Netherlands. It is the capital of the province of North Holland. Haarlem is situated at the northern edge of the Randstad, one of the most populated metropoli ...
in the Netherlands in the 1840s and 1850s. By this time, however, English gardeners had already begun to lose interest in the Proteaceae, and by the end of the 19th century European interest in the cultivation of Proteaceae was virtually non-existent. In Australia, there was little interest in the cultivation of Australian plants until the mid-20th century, despite a long-standing appreciation of their beauty as wildflowers. For example, in 1933 and 1934 '' The Western Mail'' published a series of
Edgar Dell __NOTOC__ Edgar Dell (28 November 1901 – 18 April 2008) was a Western Australian painter best known for his watercolour paintings of that state's wildflowers. Born in England in 1901, he emigrated to Western Australia in 1924, where he boug ...
paintings of Western Australian wildflowers, including a painting of ''B. sessilis''. These were subsequently republished in Charles Gardner's 1935 ''West Australian Wild Flowers''. One of the first published colour photographs of the species appeared in William Blackall's 1954 '' How to know Western Australian wildflowers'', but this publication was restricted to plant identification. The species was discussed and illustrated in the 1959 ''Wildflowers of Western Australia'', and in the 1973 ''Flowers and plants of Western Australia'', but these books did not provide cultivation advice either. Possibly the first published information on the cultivation of ''Dryandra'' appeared in the magazine ''
Australian Plants The Australian Native Plants Society (Australia) (ANPSA) is a federation of seven state-based member organisations for people interested in Australia's native flora, both in aspects of conservation and in cultivation. A national conference is h ...
'' in June and September 1961. ''D. sessilis'' was among the species treated, but as there was not yet any experimental data on cultivation, information was restricted to its aesthetic qualities and the soil in which it naturally occurs. From its inception in 1962, the
Kings Park and Botanic Garden Kings Park, (Noongar: ''Kaarta Gar-up'') is a park overlooking Perth Water and the central business district of Perth, Western Australia. The park is a mixture of grassed parkland, botanical gardens and natural bushland on Mount Eliza with ...
undertook extensive research into the cultivation of native plants, resulting in two early publications that mentioned the cultivation potential of ''B. sessilis''. In 1965,
John Stanley Beard John Stanley Beard (15 February 1916 – 17 February 2011) was a British-born forester and ecologist who resided in Australia. Beard studied at the University of Oxford where he completed his doctoral thesis on tropical forestry. While working ...
published ''Descriptive catalogue of Western Australian plants'', "a work of reference in which the horticultural characteristics of the plants concerned could be looked up by the staff", which described ''D. sessilis'' as an erect shrub with pale yellow flowers appearing from May to October, growing in sand and gravel. Five years later, Arthur Fairall published ''West Australian native plants in cultivation''. This presented largely the same information as Beard's ''catalogue'', adding only that the species flowers well in its third season.


Current knowledge

According to current knowledge, ''B. sessilis'' is an extremely hardy plant that grows in a range of soils and aspects, so long as it is given good drainage, and tolerates both drought and moderate frost. Unlike many dryandras, it grows well on limestone (alkaline) soils. It flowers very heavily and is an excellent producer of honey. It attracts birds, and is also popular with beekeepers. However, its size makes it unsuitable for smaller gardens, and if given an ideal situation it may produce a great many seedlings. It is propagated only from seed, as propagating it from cuttings has proven virtually impossible. Germination takes about five or six weeks, and plants may take two years to flower.


References


External links

* * * {{DEFAULTSORT:Banksia Sessilis sessilis Trees of Australia Ornamental trees Trees of Mediterranean climate Plants described in 1809 Eudicots of Western Australia Endemic flora of Southwest Australia