Swamp banksia | |
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Banksia paludosa flower spike Stanwell Tops, New South Wales | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Order: | Proteales |
Family: | Proteaceae |
Genus: | Banksia |
Subgenus: | Banksia subg. Banksia |
Section: | Banksia sect. Banksia |
Series: | Banksia ser. Salicinae |
Species: | B. paludosa [1] |
Binomial name | |
Banksia paludosa [1] | |
Banksia paludosa, commonly known as the marsh or swamp banksia, is a species of shrub in the plant genus Banksia . It is native to New South Wales, Australia, where it is found between Sydney and Batemans Bay, with an isolated population further south around Eden. There are two recognised subspecies, the nominate of which is a spreading shrub to 1.5 m (4.9 ft) in height, and subsp. astrolux is a taller shrub to 5 m (16 ft) high found only in Nattai National Park.
Native mammals, such as the brown antechinus and sugar glider, are important pollinators of B. paludosa. Several species of honeyeaters visit the flower spikes, as do ants and the European honey bee. The response to bushfire depends on the subspecies; subspecies paludosa regenerates from underground lignotubers, while plants of subspecies astrolux are killed by fire and regenerate from large stores of seed which have been held in cones in the plant canopy. B. paludosa is sometimes seen in cultivation, with dwarf forms being registered and sold.
The two subspecies of Banksia paludosa are identical in foliage and flower, and differ only on their size, habit, and response to bushfire. [2] Banksia paludosa subspecies paludosa is most commonly encountered as a spreading multistemmed shrub generally under 1.5 m (4.9 ft) high, or rarely 2 m (6.6 ft). In heathland habitats such as Nadgee [3] or Barren Grounds Nature Reserves, [4] it may not exceed 1 m (3.3 ft) in height. At an exposed area such as Green Cape, it is reduced further to a 30 cm (12 in) prostrate shrub. [5] This subspecies has a woody base, known as a lignotuber, which begins developing in the first year of life. [3] Banksia paludosa subspecies astrolux is a more open non-lignotuberous shrub which reaches 5 m (16 ft) high. [2]
The bark and foliage is rough and covered in multiplication signs, although the new growth is covered in fine hair. The stems are generally less than 2 cm (0.79 in) in diameter, [3] and may be red or yellow when young. [6] The leaves are alternate or whorled along the stems, and spear- to egg-shaped (lanceolate to obovate) in shape. They measure 4–13 cm (1.6–5.1 in) long and 1–3 cm (0.39–1.18 in) wide. The leaf margins are entire or have occasional serrations. The leaf undersurface is white, with a midrib. [3] Flowering occurs over autumn and winter (April to July) and the flower spikes, known as inflorescences, arise from stems that are three or more years old. Cylindrical in shape, they are composed of a central woody spike or axis from which a large number of compact floral units arise perpendicularly to it, and are generally 3.2–4 cm (1.3–1.6 in) wide and 7–13 cm (2.8–5.1 in) high. [3] The individual flowers are more openly spaced than those of other banksias, and this is especially evident in late bud. [6] This, coupled with the tall thin shape of the flower spike, makes the species quite distinctive. [3] The flower spikes are pale- to golden brown in bud, and open to a more gold colour after anthesis. [6] Variations are seen, one form having a grey limb in bud, and plants with particularly tall flower spikes have been recorded near Huskisson at Jervis Bay. [7] As with most banksias, in anthesis the opening of the individual buds proceeds up the flower spike from the base to the top (acropetal). [3] The process from bud to the finishing of flowering takes six to eight weeks. [8]
As they age, the flower spikes fade to grey, with the old flowers remaining for years. Up to 60 woody follicles develop on each spike, known in this stage as an infructescence. Narrow and elliptic, they measure 0.9–1.8 cm (0.4–0.7 in) long, 0.1–0.5 cm (0.0–0.2 in) high, and 0.3–0.7 cm (0.1–0.3 in) wide. Some follicles open spontaneously, but most remain closed until burnt by bushfire. Each follicle contains one or two fertile seeds, between which lies a woody dark brown separator of similar shape to the seeds. Measuring 1.3–1.8 cm (0.5–0.7 in) in length, the seed is obovate, and composed of a dark brown 0.8–1.3 cm (0.3–0.5 in) wide membranous "wing" and sickle-shaped (falcate) seed proper which measures 0.8–0.9 cm (0.3–0.4 in) long by 0.3–0.4 cm (0.1–0.2 in) wide. The seed surface can be smooth or covered in tiny ridges, and often glistens. The resulting seedling first grows two asymmetrical obovate cotyledon leaves measuring 0.9 cm (0.4 in) long by 0.7 cm (0.3 in) wide, which may remain for several months as several more leaves appear. The first pairs of leaves are oppositely arranged on the stem, have 3–4 "teeth" on their margins, and are narrowly obovate in shape. They are around 1.2–1.4 cm (0.5–0.6 in), and each following pair of leaves is slightly larger. [3]
The cotyledons of Banksia paludosa, B. marginata and B. integrifolia are very similar in appearance. [9] The foliages of larger shrubs of both Banksia paludosa subspecies resemble those of Banksia conferta subsp. penicillata, but the latter has a wider inflorescence, and the buds are more crowded in appearance on the inflorescence before anthesis. [10] Banksia paludosa also bears a superficial resemblance to B. oblongifolia , but the latter has a prominent midrib on the leaf underside, the new growth is covered in rusty fur, and the old spikes are bare of flowers. The latter grows on dryer rocky soils while the former grows in wetter sandy soils. [11]
Banksia paludosa was first described by Robert Brown in his 1810 On the Proteaceae of Jussieu, and named Banksia paludosa. In 1870, George Bentham demoted it to a variety of B. integrifolia (Coast Banksia), but in 1981 Alex George restored it to species rank. Its specific epithet is derived from the Latin noun palus "marsh", but is somewhat misleading, as it more often grows on sandstone ridges and heathland. Its common names, marsh banksia and swamp banksia, echo its scientific name. [12]
The current taxonomic arrangement of the genus Banksia is based on botanist Alex George's 1999 monograph for the Flora of Australia book series. [2] In this arrangement, B. paludosa is placed in Banksia subgenus Banksia , because its inflorescences take the form of Banksia's characteristic flower spikes, section Banksia because of its straight styles, and series Salicinae because its inflorescences are cylindrical. In a morphological cladistic analysis published in 1994, Kevin Thiele placed it in the newly described subseries Integrifoliae, within the series Salicinae. [13] However, this subgrouping of the Salicinae was not supported by George. [2]
B. paludosa's placement within Banksia may be summarised as follows:
Since 1998, American botanist Austin Mast and co-authors have been publishing results of ongoing cladistic analyses of DNA sequence data for the subtribe Banksiinae, which then comprised genera Banksia and Dryandra . Their analyses suggest a phylogeny that differs greatly from George's taxonomic arrangement. Banksia paludosa resolves as the closest relative, or "sister", to the three subspecies of B. integrifolia. [14] [15] [16] In 2007, Mast and Thiele rearranged the genus Banksia by merging Dryandra into it, and published B. subg. Spathulatae for the taxa having spoon-shaped cotyledons; thus B. subg. Banksia was redefined as encompassing taxa lacking spoon-shaped cotyledons. They foreshadowed publishing a full arrangement once DNA sampling of Dryandra was complete; in the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, B. paludosa is placed in B. subg. Spathulatae. [17]
Two subspecies are recognised. The more widespread subspecies paludosa is a lignotuberous shrub to 1.5 m (4.9 ft) high, while subspecies astrolux, a rare plant known only from the Starlight Trail in Nattai National Park, is a non-lignotuberous shrub which reaches 5 m (16 ft) high. [2] The latter was first recorded by contributors Brian Walters and Kevin Mills for The Banksia Atlas mapping project in the mid-1980s. They initially thought the plants were Banksia conferta subsp. penicillata until they observed the flower spikes typical of B. paludosa. The subspecies was initially termed the Nattai River form of B. paludosa, [18] until it was formally named by George, who coined its species name from the Ancient Greek aster "star" and Latin lux "light", a translation of the place it was found. [19]
Hybrids with Banksia marginata and B. integrifolia have been recorded at Nadgee Nature Reserve, where all three species occur. [5] A study of an area of extensive hybridization between B. robur and B. oblongifolia at Barren Grounds Nature Reserve revealed some plants with morphology suggestive of B. paludosa in their parentage, and requiring further investigation. [20]
Both subspecies of Banksia paludosa are endemic to New South Wales. The nominate subspecies paludosa is found from Glen Davis through to the Sydney region and then south to Ulladulla on the South Coast, with a separate population in the vicinity of Eden just north of the Victorian border. It occurs inland as far as Taralga on the Southern Tablelands. It was collected in 1966 from Hat Head on the Mid North Coast by Lawrie Johnson, but has not been found there since despite field work in the area. [5] This record aside, the northernmost historical coastal record is from what is now Centennial Park and La Perouse in Sydney's eastern suburbs, where it is now locally vanished. [8] Subspecies astrolux is restricted to Nattai National Park in the Southern Highlands. [5]
Both subspecies grow in nutrient-poor well-drained sandstone soils, in open woodland with trees such as Sydney peppermint ( Eucalyptus piperita ), silvertop ash ( E. sieberi ), grey gum ( E. punctata ), narrow-leaved stringybark ( E. sparsifolia ), red bloodwood ( Corymbia gummifera ) and smooth-barked apple ( Angophora costata ), and in heathland with species such as dwarf banksia ( Banksia oblongifolia ), coral heath ( Epacris microphylla ), and dagger hakea ( Hakea teretifolia ). [8]
Banksia paludosa subspecies paludosa is a slow-growing shrub which regenerates from bushfire by resprouting from its lignotuber. After fire, plants take around three years to flower significantly, but are flowering well by five years afterwards. Flowerhead numbers dwindle by fourteen years post bushfire. Plants are estimated to live to around 60 years of age. Seedlings also appear from seed dispersed after bushfire. [8] All banksias have developed proteoid or cluster roots in response to the nutrient-poor conditions of Australian soils (particularly lacking in phosphorus). [21]
The flower spikes of B. paludosa are unable to self-pollinate and require pollinators to set seed. [22] A 1988 isozyme study showed very high rates of outcrossing; pollen from one plant is well-mixed among other plants in the locale. Nonflying mammals are important pollinators in heathland habitat, with the brown antechinus (Antechinus stuartii) a frequent visitor to flower spikes. The sugar glider (Petaurus breviceps) is another mammal pollinator. [22] Bird species that have been observed foraging and feeding at the flowers include the red wattlebird (Anthochaera carunculata), yellow-faced honeyeater (Lichenostomus chrysops), white-eared honeyeater (L. leucotis), crescent honeyeater (Phylidonyris pyrrhoptera), New Holland honeyeater (P. novaehollandiae), and eastern spinebill (Acanthorhynchus tenuirostris). [5] Insects recorded visiting flower spikes include the European honey bee and ants. [5]
Banksia paludosa was first introduced into cultivation in England in 1805. The species was grown at Kew, Cambridge Botanic Gardens, Woburn Abbey, Loddiges nursery in Hackney, John Miller's nursery in Bristol and George Hibbert's garden at Clapham Common. [23] It was also grown in the Villa San Donato in Italy, in the collection of Anatoly Nikolaievich Demidov, 1st Prince of San Donato. [24]
B. paludosa is cultivated in Australian gardens, and does best with a sunny aspect and good drainage, in soils with a pH from 5.5 to 7.5. Slow growing, it flowers in 6 to 10 years from seed. [6] It can be propagated by seed, which take around two weeks to germinate, [25] or cutting. Low growing coastal (dwarf) forms which grow to 60 cm (2.0 ft) are also commercially available, and should be propagated by cutting to preserve features. [6] Noting the flower spikes to be "rather dull", plant author John Wrigley has described the species as "not a spectacular garden plant", [25] although its foliage has been described as "attractive". [12] A form from Jervis Bay with large orange flower spikes was deemed by amateur botanist and banksia enthusiast Alf Salkin to have horticultural potential. [26]
Banksia integrifolia, commonly known as the coast banksia, is a species of tree that grows along the east coast of Australia. One of the most widely distributed Banksia species, it occurs between Victoria and Central Queensland in a broad range of habitats, from coastal dunes to mountains. It is highly variable in form, but is most often encountered as a tree up to 25 metres (82 ft) in height. Its leaves have dark green upper surfaces and white undersides, a contrast that can be striking on windy days.
Banksia ericifolia, the heath-leaved banksia, or lantern banksia, is a species of woody shrub of the family Proteaceae native to Australia. It grows in two separate regions of Central and Northern New South Wales east of the Great Dividing Range. Well known for its orange or red autumn inflorescences, which contrast with its green fine-leaved heath-like foliage, it is a medium to large shrub that can reach 6 m (20 ft) high and wide, though is usually half that size. In exposed heathlands and coastal areas, it is more often 1–2 m (3.3–6.6 ft).
Banksia spinulosa, the hairpin banksia, is a species of woody shrub, of the genus Banksia in the family Proteaceae, native to eastern Australia. Widely distributed, it is found as an understorey plant in open dry forest or heathland from Victoria to northern Queensland, generally on sandstone though sometimes also clay soils. It generally grows as a small shrub to 2 metres (7 ft) in height, though can be a straggly tree to 6 metres (20 ft). It has long narrow leaves with inflorescences which can vary considerably in coloration; while the spikes are gold or less commonly yellowish, the emergent styles may be a wide range of colours – from black, purple, red, orange or yellow.
Banksia burdettii, commonly known as Burdett's banksia, is a species of shrub or tree of the genus Banksia in the family Proteaceae. It occurs on sandplain country north of Gingin, Western Australia. Growing to 4 m (13 ft) in height, it has long serrated leaves and large, bright flower spikes, initially white before opening to a bright orange, that appear mainly in late summer. Edmund Gilbert Baker described B. burdettii in 1934, naming it after its collector, W. Burdett.
Banksia caleyi, commonly known as Caley's banksia or red lantern banksia, is a species of woody shrub of the family Proteaceae native to Western Australia. It generally grows as a dense shrub up to 2 m (7 ft) tall, has serrated leaves and red, pendent (hanging) inflorescences which are generally hidden in the foliage. First described by Scottish naturalist Robert Brown in 1830, Banksia caleyi was named in honour of the English botanist George Caley. No subspecies are recognised. It is one of three or four related species with hanging inflorescences, which is an unusual feature within the genus.
Banksia canei, commonly known as the mountain banksia, is a species of shrub that is endemic to southeastern Australia. It is generally encountered as a many-branched shrub that grows up to 3 m (10 ft) high, with narrow leaves and the yellow inflorescences appearing from late summer to early winter. The old flowers fall off the spikes and up to 150 finely furred follicles develop, which remain closed until burnt in a bushfire. Each follicle bears two winged seeds. Response to fire is poorly known, although it is thought to regenerate by seed. Birds such as the yellow-tufted honeyeater and various insects forage among the flower spikes. It is frost tolerant in cultivation, but copes less well with aridity or humidity and is often short-lived in gardens. One cultivar, Banksia 'Celia Rosser', was registered in 1978, but has subsequently vanished.
Banksia dentata, commonly known as the tropical banksia, is a species of tree in the genus Banksia. It occurs across northern Australia, southern New Guinea and the Aru Islands. Growing as a gnarled tree to 7 m (23 ft) high, it has large green leaves up to 22 cm (8.7 in) long with dentate (toothed) margins. The cylindrical yellow inflorescences, up to 13 cm (5.1 in) high, appear over the cooler months, attracting various species of honeyeaters, sunbirds, the sugar glider and a variety of insects. Flowers fall off the ageing spikes, which swell and develop follicles containing up to two viable seeds each.
Banksia hookeriana, commonly known as Hooker's banksia, is a species of shrub of the genus Banksia in the family Proteaceae. It is native to the southwest of Western Australia and can reach up to 4 m (13 ft) high and 3 m (9.8 ft) wide. This species has long narrow serrated leaves and large, bright flower spikes, initially white before opening to a bright orange that appear over the cooler months. The flowers are pollinated by honeyeaters. The ageing flower spikes develop woody seed pods known as follicles. B. hookeriana is serotinous — large numbers of seeds are stored in the plant canopy for years until the plants are burnt by bushfire.
Banksia lemanniana, the yellow lantern banksia or Lemann's banksia, is a species of flowering plant in the family Proteaceae, native to Western Australia. It generally grows as an open woody shrub or small tree to 5 m (16 ft) high, with stiff serrated leaves and unusual hanging inflorescences. Flowering occurs over summer, the greenish buds developing into oval flower spikes before turning grey and developing the characteristic large woody follicles. It occurs within and just east of the Fitzgerald River National Park on the southern coast of the state. B. lemanniana is killed by bushfire and regenerates from seed.
Banksia marginata, commonly known as the silver banksia, is a species of tree or woody shrub in the plant genus Banksia found throughout much of southeastern Australia. It ranges from the Eyre Peninsula in South Australia to north of Armidale, New South Wales, and across Tasmania and the islands of Bass Strait. It grows in various habitats, including Eucalyptus forest, scrub, heathland and moorland. Banksia marginata varies widely in habit, ranging from a 20-centimetre (7.9 in) shrub to a 12-metre (40 ft) tree. The narrow leaves are linear and the yellow inflorescences occur from late summer to early winter. The flower spikes fade to brown and then grey and develop woody follicles bearing the winged seeds. Originally described by Antonio José Cavanilles in 1800, further collections of B. marginata were designated as several separate species by Robert Brown in 1810. However, all were reclassified as a single species by George Bentham in 1870. No distinct subspecies have been recognised by Banksia expert Alex George, who nonetheless concedes that further work is needed.
Banksia media, the southern plains banksia or golden stalk banksia, is a species of flowering plant in the family Proteaceae. An evergreen shrub, it occurs on the south coast of Western Australia between Albany and Israelite Bay, where it is a common plant. A many-branched bush with wedge-shaped serrated leaves and large golden-yellow flower spikes, known as inflorescences, it grows up to 10 metres (30 ft) high.
Banksia oblongifolia, commonly known as the fern-leaved, dwarf or rusty banksia, is a species in the plant genus Banksia. Found along the eastern coast of Australia from Wollongong, New South Wales in the south to Rockhampton, Queensland in the north, it generally grows in sandy soils in heath, open forest or swamp margins and wet areas. A many-stemmed shrub up to 3 m (9.8 ft) high, it has leathery serrated leaves and rusty-coloured new growth. The yellow flower spikes, known as inflorescences, most commonly appear in autumn and early winter. Up to 80 follicles, or seed pods, develop on the spikes after flowering. Banksia oblongifolia resprouts from its woody lignotuber after bushfires, and the seed pods open and release seed when burnt, the seed germinating and growing on burnt ground. Some plants grow between fires from seed shed spontaneously.
Banksia plagiocarpa, commonly known as the Dallachy's banksia or blue banksia, is a species of shrub or tree in the plant genus Banksia. It occurs only on Hinchinbrook Island, Queensland and the immediately adjacent mainland. First collected in 1867, Banksia plagiocarpa was not described until 1981, when Alex George named it in his monograph of the genus Banksia. Genetic studies show it to be related to Banksia aquilonia, Banksia oblongifolia and Banksia robur.
Banksia saxicola, the rock banksia or Grampians banksia, is a species of tree or shrub in the plant genus Banksia. It occurs in Victoria in two distinct populations, one in The Grampians and the other on Wilsons Promontory. Formerly considered to be a form of B. integrifolia, it was described as a distinct species by Alex George in 1981. It is most closely related to Banksia marginata.
Banksia verticillata, commonly known as granite banksia or Albany banksia, is a species of shrub or (rarely) tree of the genus Banksia in the family Proteaceae. It is native to the southwest of Western Australia and can reach up to 3 m (10 ft) in height. It can grow taller to 5 m (16 ft) in sheltered areas, and much smaller in more exposed areas. This species has elliptic green leaves and large, bright golden yellow inflorescences or flower spikes, appearing in summer and autumn. The New Holland honeyeater is the most prominent pollinator, although several other species of honeyeater, as well as bees, visit the flower spikes.
Banksia blechnifolia is a species of flowering plant in the genus Banksia found in Western Australia. It was first described by Victorian state botanist Ferdinand von Mueller in 1864, and no subspecies are recognised. It gained its specific name as its leaves are reminiscent of a fern (Blechnum). B. blechnifolia is one of several closely related species that grow as prostrate shrubs, with horizontal stems and leathery, upright leaves. The red-brown flower spikes, known as inflorescences, are up to 20 centimetres (8 in) high and appear from September to November in the Australian spring. As the spikes age, each turns grey and develops as many as 25 woody seed pods, known as follicles.
Banksia aquilonia, commonly known as the northern banksia and jingana, is a tree in the family Proteaceae and is endemic to north Queensland on Australia's northeastern coastline. With an average height of 8 m (26 ft), it has narrow glossy green leaves up to 20 cm (7.9 in) long and 6 to 10 cm high pale yellow flower spikes, known as inflorescences, appearing in autumn. As the spikes age, their flowers fall off and they develop up to 50 follicles, each of which contains two seeds.
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.
The taxonomy of Banksia integrifolia has a long and complex history, the result of confusion caused by the species' great variability, and similarities with some closely related species. The existence of hybrids between B. integrifolia and related species as well as early attempts to classify the species based on dried specimen material have also contributed to the confusion.
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. (Proteaceae), it superseded the arrangement of George Bentham, which had stood for over a hundred years. It was overturned in 1996 by Kevin Thiele and Pauline Ladiges, but restored by George in 1999. A recent publication by Austin Mast and Kevin Thiele suggests that it will soon be overturned again.