Banksia verticillata

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Granite banksia
Banksia verticillata and New Holland Honeyeater.JPG
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Proteales
Family: Proteaceae
Genus: Banksia
Species:
B. verticillata
Binomial name
Banksia verticillata
Banksia verticillata map.png
Distribution of B. verticillata in Western Australia.

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 (Phylidonyris novaehollandiae) is the most prominent pollinator, although several other species of honeyeater, as well as bees, visit the flower spikes.

Contents

A declared vulnerable species, it occurs in two disjunct populations on granite outcrops along the south coast of Western Australia, with the main population near Albany and a smaller population near Walpole, and is threatened by dieback ( Phytophthora cinnamomi ) and aerial canker ( Zythiostroma ). B. verticillata is killed by bushfire and new plants regenerate from seed afterwards. Populations take over a decade to produce seed and fire intervals of greater than twenty years are needed to allow the canopy seed bank to accumulate.

Description

Banksia verticillata grows as a spreading, bushy shrub with many branches up to 3 m (10 ft) high, but can reach 5 m (16 ft) high in sheltered locations. [2] It may be much lower or even adopt a prostrate habit in highly exposed areas which are blasted by high wind, [2] [3] or occasionally grow as a single-trunked tree. The rough grey bark has fissures, the stems and branches are finely hairy when young and become smooth with age. The leathery bright green leaves are arranged whorled, or alternately on branches, and are borne on 0.5–1.1 mm long petioles. They measure 3–9 cm (1.4–3.8 in) in length, and 0.7–1.2 cm (0.3–0.5 in) in width, and are elliptic in shape with entire (straight) recurved margins. They are initially hairy and become smooth with maturity, although their undersides remain covered with white hair. [2] The golden-yellow inflorescences appear in summer and autumn (January to April) and are 8–20 cm (3–8 in) high and 6.5 cm (2.6 in) wide. The smooth pistils are 3–3.5 cm long and hooked at the end. [4] Individual flowers open from the base of the flower spike, the wave of anthesis moving up the inflorescence. Occasionally, flowers on exposed parts may open early. It takes around 9.5 days for all flowers to open, and rates are similar during the day and night. [5] The inflorescences age to grey and the individual old flowers linger for some time before falling. Up to 100 small woody follicles may follow on old flower spikes. Measuring 1.1–1.5 cm wide, 3–4 mm wide, and jutting out 2–3 mm from the spike, they open after several years, releasing the seed. [4] Follicles more commonly appear in the middle third of the spike. The reasons for this are unknown, although timing of visits by pollinators or some anatomical factor may be relevant. [2]

Taxonomy

Discovery and naming

King George's Sound, view on the peninsula to the north of Peak Head, a field sketch executed by William Westall in December 1801. The sprawling shrub in the foreground has been tentatively identified as B. verticillata, which would make this sketch, together with another Westall sketch thought to depict B. verticillata, the earliest known drawings of the species. King George's Sound, looking north (Westall collection).jpeg
King George's Sound, view on the peninsula to the north of Peak Head, a field sketch executed by William Westall in December 1801. The sprawling shrub in the foreground has been tentatively identified as B. verticillata, which would make this sketch, together with another Westall sketch thought to depict B. verticillata, the earliest known drawings of the species.

The earliest known botanical collection of B. verticillata was made by Scottish surgeon and naturalist Archibald Menzies during the visit of the Vancouver Expedition to King George Sound in September and October 1791. As a result of this collection the species was introduced into cultivation in England, [7] yet it did not result in formal publication of the species.

The next known collection was in December 1801, during the visit of HMS Investigator to King George Sound. Little is known of the circumstances of this collection, other than what is written on the specimen label: "King Georges Sound Dec[embe]r 1801". [8] The specimen is credited to Robert Brown, but gardener Peter Good and the botanical artist Ferdinand Bauer also contributed to Brown's specimen collection, often without attribution. [9] A more precise date and location cannot be given, as neither Brown nor Good mentions the collection in his diary. [10] [11] Bauer did not publish an illustration of the species [12] and his original field sketches are lost, [13] but William Westall appears to have incorporated it into two of his field sketches, and certainly included it in the foreground of one of the oil paintings that he later worked up for the Admiralty. [6]

Brown formally described and named the species in his 1810 On the Proteaceae of Jussieu . [14] He did not identify a type specimen, but the one specimen in his collection has since been formally declared the lectotype for the species. [15] He also did not explicitly give an etymology for the specific epithet, but it is accepted that the name derives from the Latin verticillatus ("whorled"), in reference to the whorled leaf arrangement. [16]

No subspecies or varieties of Banksia verticillata have been identified and it has no taxonomic synonyms. [17] In 1891, Otto Kuntze, in his Revisio Generum Plantarum , rejected the generic name Banksia L.f., on the grounds that the name Banksia had previously been published in 1776 as Banksia J.R.Forst & G.Forst, referring to the genus now known as Pimelea . Kuntze proposed Sirmuellera as an alternative, referring to this species as Sirmuellera verticillata. [18] This application of the principle of priority was largely ignored by Kuntze's contemporaries, [19] and Banksia L.f. was formally conserved and Sirmuellera rejected in 1940. [20]

Infrageneric placement

Banksia verticillata flower spike Banksia verticillata inflorescence.jpg
Banksia verticillata flower spike

In Brown's arrangement of Banksia, B. verticillata was placed between B. compar (now B. integrifolia subsp. compar) and B. coccinea (scarlet banksia) in phyletic order. No infrageneric arrangement was provided other than the removal of one distinctive species into a subgenus of its own, because of its unusual domed flower head. As B. verticillata flowers occur in characteristic flower spikes, it was retained in Banksia verae , the "true banksias". [14] Banksia verae was renamed Eubanksia by Austrian botanist Stephan Endlicher in 1847, with B. verticillata remaining between the same two species as in Brown's sequence. [21] A more detailed arrangement was published by Carl Meissner in 1856. Eubanksia was demoted to sectional rank, and divided it into four series. B. verticillata was placed in series Salicinae because its leaves are more or less linear, and have white undersides. [22] Based as they were on leaf characters, Meissner's series were highly heterogeneous, [15] and George Bentham discarded them all in his 1870 revision of Banksia. B. verticillata was instead placed in a new section, Oncostylis , because of its hooked styles. [23] This arrangement would stand for over a century.

For many years there was confusion between B. verticillata and B. littoralis (swamp banksia). Until 1984, the latter was circumscribed as encompassing what is now Banksia seminuda (river banksia), which has whorled leaves like B. verticillata. Thus it was easy to perceive B. verticillata as falling within the range of variation of this broadly defined species. The confusion was largely cleared up once B. seminuda was recognised as a distinct taxon. [24]

Alex George published a new taxonomic arrangement of Banksia in his landmark 1981 monograph The genus Banksia L.f. (Proteaceae) . Endlicher's Eubanksia became B. subg. Banksia, and was divided into three sections, one of which was Oncostylis. Oncostylis was further divided into four series, with B. verticillata placed in series Spicigerae because its inflorescences are cylindrical. [4]

In 1996, Kevin Thiele and Pauline Ladiges published a new arrangement for the genus, after cladistic analyses yielded a cladogram significantly different from George's arrangement. With respect to B. verticillata, their findings largely accorded with George's arrangement: section Oncostylis was discarded as polyphyletic, but series Spicigerae was inferred to be monophyletic, and B. verticillata appeared in a succession of clades with the species previously identified as its closest relatives: first B. littoralis, then B. seminuda, then B. brownii, and finally B. occidentalis (red swamp banksia): [25]

B. occidentalis

B. brownii

B. seminuda

B. littoralis

B. verticillata

This clade became the basis of Thiele and Ladiges' B. subser. Occidentales, which was defined as "characterised by opposite-decussate seedling leaves and adult leaves in true whorls." [25] This arrangement stood until 1999, when George largely reverted to his 1981 arrangement in his monograph for the Flora of Australia series. Under George's taxonomic arrangement of Banksia, B. verticillata's taxonomic placement may be summarised as follows: [4]

Banksia
B. subg. Banksia
B. sect. Banksia (9 series, 50 species, 9 subspecies, 3 varieties)
B. sect. Coccinea (1 species)
B. sect. Oncostylis
B. ser. Spicigerae (7 species, 2 subspecies, 4 varieties)
B. spinulosa (4 varieties)
B. ericifolia (2 subspecies)
B. verticillata
B. seminuda
B. littoralis
B. occidentalis
B. brownii
B. ser. Tricuspidae (1 species)
B. ser. Dryandroideae (1 species)
B. ser. Abietinae (13 species, 2 subspecies, 9 varieties)
B. subg. Isostylis (3 species)

More recent molecular research by Austin Mast and colleagues provide further support of B. verticillata's placement among its nearest relatives, but these do not appear to be closely related to the remaining members of B. ser. Spicigerae, but rather occur in a clade that is sister (next closest relative) to B. nutans : [26]

B. occidentalis

B. littoralis

B. brownii

B. verticillata

B. nutans var. nutans

B. nutans var. cernuella

(B. seminuda is omitted because it was not sampled in the study, not because it occurs elsewhere in the cladogram.)

Distribution and habitat

Banksia verticillata is found in scattered populations in two disjunct segments: one clustered around Walpole, and the other around Albany and eastwards to Cheynes Beach. All but one are located within 2 km (1.5 mi) of the coast, the exception is less than 10 km (6 mi) inland. Plants grow on exposed coastal granite outcrops, often in cracks within boulders as well as shallow rocky soils. It is the only Banksia which grows exclusively in a granite soil. [27] It grows in association with Taxandria marginata , Western Australian peppermint ( Agonis flexuosa ), Andersonia sprengelioides and species of Hakea in scrub and heath. [2]

Ecology

The New Holland honeyeater (Phylidonyris novaehollandiae) is a major visitor and pollinator of Banksia verticillata. These birds can travel 15 m (50 ft) between inflorescences in a feeding session, and preferentially choose flower spikes with partly opened flowers. Other honeyeater species observed, the white-cheeked honeyeater (Phylidonyris nigra) and western spinebill (Acanthorhynchus superciliosus), visit this species to a much lesser extent. [2] The brown honeyeater (Lichmera indistincta) has also been recorded as a visitor. [28] Small mammals are not major pollinators, although bush rats (Rattus fuscipes) and house mice (Mus musculus) have been recorded. Honey bees (Apis mellifera) visit flower spikes but are not effective pollinators. [2] [5]

B. verticillata is significantly threatened by at least three microorganisms. Several populations have reduced or vanished from dieback ( Phytophthora cinnamomi ), such as those at Two Peoples Bay Nature Reserve and Gull Rock National Park. The honey fungus Armillaria luteobubalina has killed plants in Torndirrup National Park, and aerial canker ( Zythiostroma ) has decimated populations at Waychinicup National Park east of Albany. [2]

B. verticillata plants are generally killed by fire and regenerate from seed. A field study after a mild fire in Torndirrup National Park published in 1994 found that plants burnt by fire were ten times as likely to have seedlings come up under their crown as unburnt plants (with an average of 25.2 seedlings per burnt plant), and burnt spikes released double the number of seeds as unburnt spikes. Despite this, interfire recruitment (seedlings arising between fires) has also been recorded, and might be more common than in other Banksia species. Observations at several of the populations showed many plants produced their first seed anywhere from 13 to 17 years of age, leading to a recommendation of 20 years between fires to allow seed banks to accumulate. [2] If fire occurs too frequently, plants are burned before reaching maturity or before they have produced sufficient seed to ensure regeneration of the population. This may cause a population decline or even local extinction. Too long a time between fires also causes population decline, as more plants die of natural attrition without releasing their seed, resulting in seed wastage. [29]

Conservation

Banksia verticillata growing in a garden in Albany Banksia verticillata habit.jpg
Banksia verticillata growing in a garden in Albany

Banksia verticillata has been declared vulnerable under the federal Environment Protection and Biodiversity Conservation Act 1999 , [1] and Rare under Western Australia's Wildlife Conservation Act 1950 . [30] A 1995 census yielded an estimated total population size of 4500 plants. [2] Apart from dieback and aerial canker, frequency of bushfires and illegal picking have been cited as threats. Ongoing management includes monitoring of current populations, gathering more data on best response to fire, and restricting access to populations. [31] Seed has been collected from many populations, but germination rates after ten years of storage are much lower than in Banksia brownii . [2] Translocation is considered an option in the future, [31] as is spraying with phosphite, particularly in the vicinity of Walpole. [2] Used successfully on B. brownii but as yet untrialled with B. verticillata, phosphite boosts the resistance of both infected and uninfected plants, and also acts as a direct fungicide. Aerial spraying of phosphite boosts plant survival and slows the spread of infection, [32] [33] but must be carefully managed as studies have shown that foliar spraying of phosphite adversely affects root and shoot growth. [34]

Cultivation

Banksia verticillata is seldom seen in cultivation. The natural growing conditions point to a sunny aspect and good drainage as being important in cultivation. It is good for coastal situations and erosion control. [35] Very sensitive to dieback, B. verticillata (like most other western Australian banksias) perishes quickly in humid conditions or poor drainage. It has been grafted successfully onto Banksia integrifolia . [36] Seeds do not require any treatment, and take 19 to 49 days to germinate. [37]

Related Research Articles

<i>Banksia</i> Genus of flowering plants in the family Proteaceae

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. Banksias range in size from prostrate woody shrubs to trees up to 30 metres tall. They are found in a wide variety of landscapes: sclerophyll forest, (occasionally) rainforest, shrubland, and some more arid landscapes, though not in Australia's deserts.

<i>Banksia coccinea</i> Species of shrub or small tree

Banksia coccinea, commonly known as the scarlet banksia, waratah banksia or Albany banksia, is an erect shrub or small tree in the family Proteaceae. Its distribution in the wild is along the south west coast of Western Australia, from Denmark to the Stokes National Park, and north to the Stirling Range, growing on white or grey sand in shrubland, heath or open woodland. Reaching up to 8 m (26 ft) in height, it is a single-stemmed plant that has oblong leaves, which are 3–9 cm (1.2–3.5 in) long and 2–7 cm (0.8–2.8 in) wide. The prominent red and white flower spikes appear mainly in the spring. As they age they develop small follicles that store seeds until opened by fire. Though widely occurring, it is highly sensitive to dieback and large populations of plants have succumbed to the disease.

<i>Banksia brownii</i> Species of shrub in the family Proteaceae found in Australia

Banksia brownii, commonly known as feather-leaved banksia or Brown's banksia, is a species of shrub that grows in southwest Western Australia. A plant with fine feathery leaves and large red-brown flower spikes, it usually grows as an upright bush around two metres (6.6 ft) high, but can also occur as a small tree or a low spreading shrub. First collected in 1829 and published the following year, it is placed in Banksiasubgenus Banksia, section Oncostylis, series Spicigerae. There are two genetically distinct forms.

<i>Banksia ericifolia</i> Woody shrub native to Australia

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).

<i>Banksia menziesii</i> Species of plant

Banksia menziesii, commonly known as firewood banksia, is a species of flowering plant in the genus Banksia. It is a gnarled tree up to 10 m (33 ft) tall, or a lower spreading 1–3 m (3.3–9.8 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 inflorescences are often two-coloured red or pink and yellow, and their colour has given rise to more unusual common names such as port wine banksia and strawberry banksia. Yellow blooms are rarely seen.

Taxonomy of <i>Banksia</i> Classification of the plant genus Banksia

As with other flowering plants, the taxonomy of Banksia has traditionally been based on anatomical and morphological properties of the Banksia flower, fruiting structure and seed, along with secondary characteristics such as leaf structure and growth habit. Increasingly, molecular evidence from DNA is providing important new insights into relationships within the genus and between this and other genera in the Proteaceae.

<i>Banksia canei</i> Shrub species endemic to Australia

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.

<i>Banksia lemanniana</i> Shrub of the family Proteaceae native to Western Australia

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.

<i>Banksia nutans</i> Species of shrub in the family Proteaceae native to the south coast of Western Australia

Banksia nutans, commonly known as nodding banksia, is a species of shrub native to the south coast of Western Australia in the genus Banksia. Growing to a metre (3 ft) tall, it has pale blue-green fine-leaved foliage and unusual purple-brown inflorescences which hang upside down rather than grow upright like most other banksias.

<i>Banksia seminuda</i> Species of tree in the family Proteaceae found in south west Western Australia

Banksia seminuda, commonly known as the river banksia, is a tree in the plant genus Banksia. It is found in south west Western Australia from Dwellingup (32°42′ S) to the Broke Inlet east of Denmark (34°57′ S). It is often mistaken for, and was originally considered a subspecies of, the Banksia littoralis. Stephen Hopper described the subspecies remanens as a short-leaved shrubby form found in the coastal sands below granite outcrops in the Walpole-Nornalup National Park, however George does not feel this form warrants taxonomic recognition as it lies within the normal variability of the species and there was no clear distinction between it and the other populations of B. seminuda.

<i>Banksia telmatiaea</i> Australian shrub that grows in marshes and swamps

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 metres tall, with narrow leaves and a pale brown flower spike, which can produce profuse quantities of nectar. First collected in the 1840s, it was not published as a separate species until 1981; as with several other similar species it was previously included in B. sphaerocarpa.

<i>Banksia violacea</i> Species of plant

Banksia violacea, commonly known as violet banksia, is a species of shrub or tree in the plant genus Banksia. It generally grows as a small shrub to 1.5 m (5 ft) high with fine narrow leaves, and is best known for its unusually coloured dark purple-violet inflorescences. The colour of the inflorescences, short leaves, and flattened follicles which are sticky when young, help identify this species from others in the field. It is found in low shrubland in southern regions of Western Australia from Esperance in the east to Narrogin in the west, growing exclusively in sandy soils.

The ecology of Banksia is the relationships and interactions among the plant genus Banksia and its environment. Banksia has a number of adaptations that have so far enabled the genus to survive despite dry, nutrient-poor soil, low rates of seed set, high rates of seed predation and low rates of seedling survival. These adaptations include proteoid roots and lignotubers; specialised floral structures that attract nectariferous animals and ensure effective pollen transfer; and the release of seed in response to bushfire.

<i>Banksia <span style="font-style:normal;">ser.</span> Spicigerae</i> Taxonomic series in the family Proteaceae

Banksia ser. Spicigerae is a taxonomic series in the genus Banksia. It consists of the seven species in section Oncostylis that have cylindrical inflorescences. These range in form from small shrubs to tall trees. The leaves grow in either an alternate or whorled pattern, with various shape forms. The Spicigerae inflorescence is held erect, subtended by a whorl of branchlets, and retains a regular pattern until anthesis. The perianth limb is horizontal until anthesis, at which point the perianth opens from underneath. The pollen-presenter is ovoid or conical. The seed wings are not notched.

<span class="mw-page-title-main">Bentham's taxonomic arrangement of Banksia</span> 1870 arrangement of the Australian endemic plant genus 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 monograph The genus Banksia L.f. (Proteaceae).

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. It replaced Alex George's 1981 arrangement, but most aspects were not accepted by George, and it was soon replaced by a 1999 revision of George's arrangement. However some herbaria have continued to follow Thiele and Ladiges on some points.

<span class="mw-page-title-main">George's taxonomic arrangement of Banksia</span> 1981 and 1996 arrangements of the Australian endemic plant genus 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. (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.

<i>Banksia spinulosa <span style="font-style:normal;">var.</span> collina</i> Variety of shrub in the family Proteaceae from the east coast of Australia

Banksia spinulosa var. collina is a shrub that grows along the east coast of Australia, in Queensland and New South Wales. Commonly known as Hill Banksia or Golden Candlesticks, it is a taxonomic variety of B. spinulosa. It is a popular garden plant widely sold in nurseries.

<i>Banksia spinulosa <span style="font-style:normal;">var.</span> spinulosa</i> Variety of shrub in the family Proteaceae from the east coast of Australia

Banksia spinulosa var. spinulosa is a shrub that grows along the east coast of Australia, in Queensland and New South Wales.

<i>Banksia spinulosa <span style="font-style:normal;">var.</span> cunninghamii</i> Variety of shrub in the family Proteaceae from the east coast of Australia

Banksia spinulosa var. cunninghamii, sometimes given species rank as Banksia cunninghamii, is a shrub that grows along the east coast of Australia, in Victoria and New South Wales. It is a fast-growing non-lignotuberous shrub or small tree infrequently cultivated.

References

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