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:::::''[[Banksia aculeata|B. aculeata]]''
:::::''[[Banksia aculeata|B. aculeata]]''


Since 1998, [[Austin Mast]] has been publishing results of ongoing cladistic analyses of [[DNA sequence]] data for the subtribe [[Banksiinae]]. His analyses suggest a [[phylogeny]] that is rather different from previous taxonomic arrangements, but support the placement of ''B.&nbsp;aculeata'' alongside ''B.&nbsp;caleyi'' in a clade corresponding closely with ''B.''&nbsp;ser. ''Tetragonae''.<ref name="Mast 1998">{{cite journal | last = Mast | first=Austin R. | author-link=Austin Mast | year = 1998 | title = Molecular Systematics of Subtribe Banksiinae (''Banksia'' and ''Dryandra''; Proteaceae) Based on cpDNA and nrDNA Sequence Data: Implications for Taxonomy and Biogeography | journal = Australian Systematic Botany | volume = 11 | issue = 4 | pages = 321–42 | doi = 10.1071/SB97026}}</ref><ref name="Mast 2002">{{cite journal | last1 = Mast | first1 = Austin R. | last2=Givnish | first2=Thomas J. | author-link2 = Thomas J. Givnish | year = 2002 | title = Historical Biogeography and the Origin of Stomatal Distributions in ''Banksia'' and ''Dryandra'' (Proteaceae) based on Their cpDNA Phylogeny | journal = [[American Journal of Botany]] | volume = 89 | issue = 8 | issn = 0002-9122 | pages = 1311–1323 | doi = 10.3732/ajb.89.8.1311 | pmid=21665734| s2cid = 2835600 | url = http://pdfs.semanticscholar.org/8281/1c898a77b2a26b0a2aa25c69dd9269d1d3e0.pdf | archive-url = https://web.archive.org/web/20190227234058/http://pdfs.semanticscholar.org/8281/1c898a77b2a26b0a2aa25c69dd9269d1d3e0.pdf | url-status = dead | archive-date = 2019-02-27 | doi-access = free }}</ref><ref name="Mast 2005">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Jones | first2 = Eric H. | last3 = Havery | first3 = Shawn P. | year = 2005 | volume = 18 | issue = 1 | title = An Assessment of Old and New DNA Sequence Evidence for the Paraphyly of ''Banksia'' with Respect to ''Dryandra'' (Proteaceae) | journal = Australian Systematic Botany | pages = 75–88 | doi = 10.1071/SB04015}}</ref>
Since 1998, [[Austin Mast]] has been publishing results of ongoing cladistic analyses of [[DNA sequence]] data for the subtribe [[Banksiinae]]. His analyses suggest a [[phylogeny]] that is rather different from previous taxonomic arrangements, but support the placement of ''B.&nbsp;aculeata'' alongside ''B.&nbsp;caleyi'' in a clade corresponding closely with ''B.''&nbsp;ser. ''Tetragonae''.<ref name="Mast 1998">{{cite journal | last = Mast | first=Austin R. | author-link=Austin Mast | year = 1998 | title = Molecular Systematics of Subtribe Banksiinae (''Banksia'' and ''Dryandra''; Proteaceae) Based on cpDNA and nrDNA Sequence Data: Implications for Taxonomy and Biogeography | journal = Australian Systematic Botany | volume = 11 | issue = 4 | pages = 321–42 | doi = 10.1071/SB97026}}</ref><ref name="Mast 2002">{{cite journal | last1 = Mast | first1 = Austin R. | last2=Givnish | first2=Thomas J. | author-link2 = Thomas J. Givnish | year = 2002 | title = Historical Biogeography and the Origin of Stomatal Distributions in ''Banksia'' and ''Dryandra'' (Proteaceae) based on Their cpDNA Phylogeny | journal = [[American Journal of Botany]] | volume = 89 | issue = 8 | issn = 0002-9122 | pages = 1311–1323 | doi = 10.3732/ajb.89.8.1311 | pmid=21665734| s2cid = 2835600 | doi-access = free }}</ref><ref name="Mast 2005">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Jones | first2 = Eric H. | last3 = Havery | first3 = Shawn P. | year = 2005 | volume = 18 | issue = 1 | title = An Assessment of Old and New DNA Sequence Evidence for the Paraphyly of ''Banksia'' with Respect to ''Dryandra'' (Proteaceae) | journal = Australian Systematic Botany | pages = 75–88 | doi = 10.1071/SB04015}}</ref>


Early in 2007, Mast and Thiele initiated a rearrangement by transferring ''Dryandra'' to ''Banksia'', and publishing ''B.''&nbsp;subg. ''Spathulatae'' for the species having spoon-shaped cotyledons; in this way, they also redefined the [[autonym (botany)|autonym]] ''B.''&nbsp;subg. ''Banksia''. They have refrained from publishing a full arrangement of ''Banksia'' until DNA sampling of ''Dryandra'' is complete; in the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then ''B.&nbsp;caleyi'' is placed in ''B.''&nbsp;subg. ''Banksia''.<ref name="Mast 2007">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Thiele | first2 = Kevin | year = 2007 | title = The Transfer of ''Dryandra'' R.Br. to ''Banksia'' L.f. (Proteaceae) | journal = [[Australian Systematic Botany]] | volume = 20 | pages = 63–71 | doi = 10.1071/SB06016}}</ref>
Early in 2007, Mast and Thiele initiated a rearrangement by transferring ''Dryandra'' to ''Banksia'', and publishing ''B.''&nbsp;subg. ''Spathulatae'' for the species having spoon-shaped cotyledons; in this way, they also redefined the [[autonym (botany)|autonym]] ''B.''&nbsp;subg. ''Banksia''. They have refrained from publishing a full arrangement of ''Banksia'' until DNA sampling of ''Dryandra'' is complete; in the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then ''B.&nbsp;caleyi'' is placed in ''B.''&nbsp;subg. ''Banksia''.<ref name="Mast 2007">{{cite journal | last1 = Mast | first1 = Austin R. | last2 = Thiele | first2 = Kevin | year = 2007 | title = The Transfer of ''Dryandra'' R.Br. to ''Banksia'' L.f. (Proteaceae) | journal = [[Australian Systematic Botany]] | volume = 20 | pages = 63–71 | doi = 10.1071/SB06016}}</ref>

Revision as of 10:40, 8 May 2021

Banksia caleyi
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Proteales
Family: Proteaceae
Genus: Banksia
Species:
B. caleyi
Binomial name
Banksia caleyi
Synonyms[1]
  • Banksia caleyi R.Br. var. caleyi
  • Banksia caleyi var. sinuosa Meisn.
  • Sirmuellera caleyi (R.Br.) Kuntze

Banksia caleyi, commonly known as Caley's banksia[2] 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.

Found south and east of the Stirling Ranges through to the vicinity of Jerramungup, Banksia caleyi grows in a habitat marked by periodic bushfires. Plants are killed by fire and regenerate by seed afterwards. The species was classified as "Not Threatened" under the Wildlife Conservation Act of Western Australia. In contrast to most other Western Australian banksias, it appears to have some resistance to dieback from the soil-borne water mould Phytophthora cinnamomi, and is comparatively easy to grow in cultivation.

Description

Cream and pink flowers

Banksia caleyi grows as a many-branched bushy shrub to 2 m (7 ft) in height, with crumbly grey bark.[3] Rarely, plants of up to 4 m (13 ft) have been found.[4] The new growth is hairy,[3] and generally occurs in summer.[4] The branchlets become smooth after around two years. The stiff leaves are narrowly wedge-shaped (cuneate) and measure 5–14 cm (2–5+1⁄2 in) in length by 1.3–2.4 cm (1⁄2–1 in) wide. The leaf margins are serrated, with many teeth measuring 0.4–0.6 cm (1⁄8–1⁄4 in) each.[3]

Flowering takes place between September and January.[4] The inflorescences hang down from the ends of three- to five-year-old branchlets deep within the shrub and measure 5–9 cm (2–3+1⁄2 in) in length and roughly 7 cm (2+3⁄4 in) in diameter. The flowers are cream at the base and deep pink to red in the upper half, and are brightest before anthesis and then gradually fade with age.[3] The inflorescences eventually turn grey, the old flowers remaining as up to 25 large woody follicles develop. Oval in shape and covered with fine hair, the follicles can reach 4 cm (1+5⁄8 in) long, 2.5 cm (1 in) high, and 2.5 cm (1 in) wide.[5]

The obovate seed is 4.3–4.7 cm (1+3⁄4–1+7⁄8 in) long and fairly flattened, and is composed of the wedge-shaped seed body proper, measuring 1.4–1.5 cm (1⁄2–5⁄8 in) long and 1.6–1.7 cm (5⁄8 in) wide, and a papery wing. One side, termed the outer surface, is dark brown and wrinkled, while the other is black and smooth. Both surfaces sparkle slightly. The seeds are separated by a sturdy dark brown seed separator that is roughly the same shape as the seeds with a depression where the seed body sits adjacent to it in the follicle. Seedlings have cuneate cotyledons which measure 1.1–1.3 cm (3⁄8–1⁄2 in) long and 1.3–1.4 cm (1⁄2 in) wide. These are dull green with three veins, and the margin of the wedge may be red and crenulated (lined with small teeth). The hypocotyl is red and measures 1.5–2 cm (5⁄8–3⁄4 in) high. Seedlings have hairy stems and leaves that are oppositely arranged (arising from the stem in pairs) that are obovate with triangular-lobed serrate margins.[3]

Taxonomy

Main article: Taxonomy of Banksia

Robert Brown formally described Banksia caleyi in his 1830 work Supplementum primum Prodromi florae Novae Hollandiae,[6] naming it in honour of the English botanist George Caley. The type specimen was collected by William Baxter, inland from King George Sound on Western Australia's south coast, in 1829.[7]

Carl Meissner placed B. caleyi in series Quercinae in his 1856 arrangement of the genus on account of its strongly dentate, cuneate to obovate leaves.[8] As they were defined on leaf characters alone, all of Meissner's series were highly heterogeneous. Meissner also described B. caleyi variety sinuosa from material collected by James Drummond,[8] which was reviewed by Alex George and found to be no different from the other collections of B. caleyi. Drummond also collected material identified as B. caleyi that was named as a distinct species—Banksia aculeata—in 1981.[3] No subspecies of B. caleyi itself are recognised.[5]

George Bentham published a thorough revision of Banksia in his landmark publication Flora Australiensis in 1870. In Bentham's arrangement, the number of recognised Banksia species was reduced from 60 to 46. Bentham defined four sections based on leaf, style and pollen-presenter characters. Banksia caleyi was placed in section Orthostylis.[9] 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 caleyi.[10] This application of the principle of priority was largely ignored by Kuntze's contemporaries,[11] and Banksia L.f. was formally conserved and Sirmuellera rejected in 1940.[12]

In his 1981 monograph The genus Banksia L.f. (Proteaceae), Alex George placed B. caleyi in B. subg. Banksia because its inflorescence is a typical Banksia flower spike shape; in B. sect. Banksia because of its straight styles; and B. ser. Tetragonae because of its pendulous inflorescences. He considered its closest relative to be B. aculeata, which has narrower leaves with fewer, larger lobes; longer perianths, which grade from red to cream rather than from cream to red; shorter pistils; and also differences in the follicles, seeds and flowering time.[3]

In 1996, Kevin Thiele and Pauline Ladiges published the results of a cladistic analysis of morphological characters of Banksia. They retained George's subgenera and many of his series, but discarded his sections. George's B. ser. Tetragonae was found to be monophyletic, and therefore retained; and their analysis of the relationships within the series supported the placement of B. caleyi alongside B. aculeata.[13]

Old flower spike showing large follicles, Muséum de Toulouse
Shrub in cultivation at Mount Barker, Western Australia

B. caleyi's placement in Thiele and Ladiges' arrangement may be summarised as follows:[13]

Banksia
B. subg. Isostylis (3 species)
B. elegans (incertae sedis)
B. subg. Banksia
B. ser. Tetragonae
B. elderiana
B. lemanniana
B. caleyi
B. aculeata

The arrangement of Thiele and Ladiges was not accepted by George, and was discarded in his 1999 revision. Under George's 1999 arrangement, B. caleyi's placement was as follows:[5]

Banksia
B. subg. Banksia
B. sect. Banksia
B. ser. Salicinae (11 species, 7 subspecies)
B. ser. Grandes (2 species)
B. ser. Banksia (8 species)
B. ser. Crocinae (4 species)
B. ser. Prostratae (6 species, 3 varieties)
B. ser. Cyrtostylis (13 species)
B. ser. Tetragonae
B. lemanniana
B. caleyi
B. aculeata

Since 1998, Austin Mast has been publishing results of ongoing cladistic analyses of DNA sequence data for the subtribe Banksiinae. His analyses suggest a phylogeny that is rather different from previous taxonomic arrangements, but support the placement of B. aculeata alongside B. caleyi in a clade corresponding closely with B. ser. Tetragonae.[14][15][16]

Early in 2007, Mast and Thiele initiated a rearrangement by transferring Dryandra to Banksia, and publishing B. subg. Spathulatae for the species having spoon-shaped cotyledons; in this way, they also redefined the autonym B. subg. Banksia. They have refrained from publishing a full arrangement of Banksia until DNA sampling of Dryandra is complete; in the meantime, if Mast and Thiele's nomenclatural changes are taken as an interim arrangement, then B. caleyi is placed in B. subg. Banksia.[17]

Distribution and habitat

Distribution in southern Western Australia

Banksia caleyi is found near the southern coast of Western Australia, from South Stirling to the West River and northeast to Pingrup. Some of its population lies within Fitzgerald River National Park.[4] Often locally abundant,[3] it is found in mallee woodland on white sand, gravel, and sandy clay, generally on flat or slightly undulating land.[4] The annual rainfall is 550–600 mm (22–24 in).[7] Banksia caleyi is classified as Not Threatened under the Wildlife Conservation Act of Western Australia.[2]

Ecology

See also: Ecology of Banksia

Like many plants in south-west Western Australia, Banksia caleyi is adapted to an environment in which bushfire events are relatively frequent. Most Banksia species can be placed in one of two broad groups according to their response to fire: Reseeders are killed by fire, but fire triggers the release of their canopy seed bank, thus promoting recruitment of the next generation. Resprouters survive fire, resprouting from a lignotuber or, more rarely, epicormic buds protected by thick bark.[18] B. caleyi belongs to the reseeder group. In the wild, seedlings take at least three to four years to reach flowering after bushfire.[4] Non-patchy fires occurring at intervals of less than seven years may wipe out local populations of reseeders.[18]

Banksia caleyi has been shown to have a low susceptibility to dieback from the soil-borne water mould Phytophthora cinnamomi, unlike most Western Australian banksias.[19] The fungal pathogen Botryosphaeria ribis has been recovered from B. caleyi.[20] The caterpillar of the dryandra moth (Carthaea saturnioides) feeds on the leaves, though it prefers to eat those of dryandra species that grow alongside it.[21]

The upside-down flower spikes drip nectar onto the ground or lower leaves, suggesting pollination by nonflying mammals which are attracted to the scent.[22] Supporting this hypothesis, the spiky leaves seem to also prevent access to foragers not at ground level.[3] Furthermore, the individual flower structure is similar to Banksia attenuata, for which the honey possum (Tarsipes rostratus) is a major pollinator.[22]

Cultivation

Seeds do not require any treatment, and take 23 to 50 days to germinate in cultivation.[23] Banksia caleyi is a medium- slow-growing plant, taking four to five years to flower from seed.[7] The flowers are attractive but are obscured by the foliage. This species can grow in a range of soil types so long as they provide good drainage. The nominal soil pH range is from 6 to 7.5. It grows in full sun and partly shaded situations, and tolerates light pruning. Unlike many other Western Australian banksias, Banksia caleyi has had some degree of success in growing in more humid areas, such as Australia's east coast. It attracts pygmy and honey possums in the garden.[7]

References

  1. ^ a b "Banksia caleyi". Australian Plant Census. Retrieved 10 April 2020.
  2. ^ a b "Banksia caleyi R.Br". FloraBase. Western Australian Government Department of Biodiversity, Conservation and Attractions.
  3. ^ a b c d e f g h i George, Alex S. (1981). "The Genus Banksia L.f. (Proteaceae)". Nuytsia. 3 (3): 239–473 [383–86]. ISSN 0085-4417.
  4. ^ a b c d e f Taylor, Anne; Hopper, Stephen (1988). The Banksia Atlas (Australian Flora and Fauna Series Number 8). Canberra: Australian Government Publishing Service. ISBN 0-644-07124-9. pp. 70–71.
  5. ^ a b c George, Alex (1999). "Banksia". In Wilson, Annette (ed.). Flora of Australia: Volume 17B: Proteaceae 3: Hakea to Dryandra. Collingwood, Victoria: CSIRO Publishing / Australian Biological Resources Study. pp. 175–251. ISBN 0-643-06454-0.
  6. ^ Brown, Robert (1830). Supplementum Primum Prodromi Florae Novae Hollandiae. London, United Kingdom: Richard Taylor. p. 35.
  7. ^ a b c d Collins, Kevin; Collins, Kathy; George, Alex (2008). Banksias. Melbourne: Bloomings Books Pty Ltd. p. 169. ISBN 978-1-876473-58-7.
  8. ^ a b Meissner, Carl (1856). "Proteaceae". In de Candolle, A. P. (ed.). Prodromus systematis naturalis regni vegetabilis, sive, Enumeratio contracta ordinum generum specierumque plantarum huc usque cognitarium, juxta methodi naturalis, normas digesta. Prodromus Systematis Naturalis Regni Vegetabilis. Vol. 14. Paris, France: Sumptibus Sociorum Treuttel et Wurtz. p. 462.
  9. ^ Bentham, George (1870). "Banksia" . Flora Australiensis: Volume 5: Myoporineae to Proteaceae. London, United Kingdom: L. Reeve & Co. pp. 541–62.
  10. ^ Kuntze, Otto (1891). Revisio generum plantarum. Vol. 2. Leipzig: Arthur Felix. pp. 581–582.
  11. ^ Rehder, A.; Weatherby, C. A.; Mansfeld, R.; Green, M. L. (1935). "Conservation of Later Generic Homonyms". Bulletin of Miscellaneous Information (Royal Botanic Gardens, Kew). 1935 (6/9): 368. doi:10.2307/4107078. JSTOR 4107078.
  12. ^ Sprague, T. A. (1940). "Additional Nomina Generica Conservanda (Pteridophyta and Phanerogamae)". Royal Botanic Gardens, Kew. 1940 (3): 99. doi:10.2307/4111642. JSTOR 4111642.
  13. ^ a b Thiele, Kevin; Ladiges, Pauline Y. (1996). "A Cladistic Analysis of Banksia (Proteaceae)". Australian Systematic Botany. 9 (5): 661–733. doi:10.1071/SB9960661.
  14. ^ Mast, Austin R. (1998). "Molecular Systematics of Subtribe Banksiinae (Banksia and Dryandra; Proteaceae) Based on cpDNA and nrDNA Sequence Data: Implications for Taxonomy and Biogeography". Australian Systematic Botany. 11 (4): 321–42. doi:10.1071/SB97026.
  15. ^ Mast, Austin R.; Givnish, Thomas J. (2002). "Historical Biogeography and the Origin of Stomatal Distributions in Banksia and Dryandra (Proteaceae) based on Their cpDNA Phylogeny". American Journal of Botany. 89 (8): 1311–1323. doi:10.3732/ajb.89.8.1311. ISSN 0002-9122. PMID 21665734. S2CID 2835600.
  16. ^ Mast, Austin R.; Jones, Eric H.; Havery, Shawn P. (2005). "An Assessment of Old and New DNA Sequence Evidence for the Paraphyly of Banksia with Respect to Dryandra (Proteaceae)". Australian Systematic Botany. 18 (1): 75–88. doi:10.1071/SB04015.
  17. ^ Mast, Austin R.; Thiele, Kevin (2007). "The Transfer of Dryandra R.Br. to Banksia L.f. (Proteaceae)". Australian Systematic Botany. 20: 63–71. doi:10.1071/SB06016.
  18. ^ a b Lamont, Byron B.; Markey, Adrienne (1995). "Biogeography of Fire-killed and Resprouting Banksia Species in South-western Australia". Australian Journal of Botany. 43 (3): 283–303. doi:10.1071/BT9950283.
  19. ^ McCredie, Thomas A.; Dixon, Kingsley W.; Sivasithamparam, Krishnapillai (1985). "Variability in the Resistance of Banksia L.f. Species to Phytophthora cinnamomi Rands". Australian Journal of Botany. 33 (6): 629–37. doi:10.1071/BT9850629.
  20. ^ Zhou, Shiguo; Stanosz, Glen R. (2001). "Relationships Among Botryosphaeria Species and Associated Anamorphic Fungi Inferred from the Analyses of ITS and 5.8S rDNA Sequences". Mycologia. 93 (3): 516–27. doi:10.2307/3761737. JSTOR 3761737.
  21. ^ McFarland, Noel (1979). "Annotated List of Larval Foodplant Records for 280 Species of Australian Moths". Journal of the Lepidopterists' Society. 33 (supplement): 1–72 [19].
  22. ^ a b Weins, Delbert; Renfree, Marilyn; Wooller, Ronald D. (1979). "Pollen Loads of Honey Possums (Tarsipes spencerae) and Non-flying Mammal Pollination in South-western Australia". Annals of the Missouri Botanical Garden. 66 (4): 830–38. doi:10.2307/2398921. JSTOR 2398921.
  23. ^ Sweedman, Luke; Merritt, David, eds. (2006). Australian Seeds: a Guide to their Collection, Identification and Biology. Collingwood, Victoria: CSIRO Publishing. p. 202. ISBN 0-643-09298-6.
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