Banksia telmatiaea
Encyclopedia
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
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

. It grows as an upright bush up to 2 m (7 ft) 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 species until 1981; as with several other similar species it was previously included in B. sphaerocarpa
Banksia sphaerocarpa
Banksia sphaerocarpa, commonly known as the Fox Banksia or Round-fruit Banksia, is a species of shrub or tree in the plant genus Banksia . It is generally encountered as a 1–2 m high shrub, and is usually smaller in the north of its range...

(Fox Banksia).

The shrub grows amongst scrubland in seasonally wet lowland areas of the coastal sandplain between Badgingarra
Badgingarra, Western Australia
Badgingarra is a small town located in the Wheatbelt region of Western Australia, about north of Perth in the Shire of Dandaragan.It lies on the Brand Highway adjacent to the Badgingarra National Park.-History:...

 and Serpentine
Serpentine, Western Australia
Serpentine is a town located south-southeast of Perth, the capital of Western Australia, and 7 km south of Mundijong. Serpentine is located on the South Western Railway between Perth and Bunbury, and was one of the original stations when the line was opened in 1893...

 in Western Australia
Western Australia
Western Australia is a state of Australia, occupying the entire western third of the Australian continent. It is bounded by the Indian Ocean to the north and west, the Great Australian Bight and Indian Ocean to the south, the Northern Territory to the north-east and South Australia to the south-east...

. A little studied species, not much is known of its ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

 or conservation biology
Conservation biology
Conservation biology is the scientific study of the nature and status of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction...

. Reports do suggest, however, that it is pollinated by a variety of birds and small mammals. Like many members of series Abietinae
Banksia ser. Abietinae
Banksia ser. Abietinae is avalid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

, it has not been considered to have much horticultural potential and is rarely cultivated.

Description

B. telmatiaea grows as an upright bush up to 2 m (7 ft) high. It has hairy stems and branchlets, and straight, narrow leaves from 1½ to 3 cm (½–1 in) long and about a millimetre ( in) wide.
Flowers occur in "flower spikes", inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

s made up of hundreds of flower pairs densely packed around a woody axis. B. telmatiaea's inflorescences are roughly spherical, with a diameter of 3–5 cm (1–2 in). It contains between 500 and 900 golden brown to pale brown flowers, each of which consists of a tubular perianth
Perianth
The term perianth has two similar but separate meanings in botany:* In flowering plants, the perianth are the outer, sterile whorls of a flower...

 made up of four fused tepal
Tepal
Tepals are elements of the perianth, or outer part of a flower, which include the petals or sepals. The term tepal is more often applied specifically when all segments of the perianth are of similar shape and color, or undifferentiated, which is called perigone...

s, and one long wiry style. The styles are hooked rather than straight, and are initially trapped inside the upper perianth parts, but break free at anthesis
Anthesis
Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period.The onset of anthesis is spectacular in some species. In Banksia species, for example, anthesis involves the extension of the style far beyond the upper perianth parts...

. The species generally flowers from April to August, although flowers have been observed as late as November. They take five to six weeks to develop from bud, then reach anthesis over a period of two weeks. The flowers produce unusually large quantities of nectar; indeed some flowers produce so much that it drips to the ground.

The fruiting structure is a stout woody "cone", with a hairy appearance caused by the persistence of old withered flower parts. This may be embedded with up to 70 woody follicles
Follicle (fruit)
In botany, a follicle is a dry unilocular many-seeded fruit formed from one carpel and dehiscing by the ventral suture in order to release seeds, such as in larkspur, magnolia, banksia, peony and milkweed....

, each of which contains a single seed. As with other Banksia species, only a small proportion of flowers go on to form follicles; in the case of B. telmatiaea, the proportion is around 4% for those "cones" that set some fruit. However, about 80% of fruiting structures set no fruit at all. According to John K. Scott, "there [is] no obvious reason on the basis of morphology of pollination for this lack of seed set".

Taxonomy


Discovery and naming

B. telmatiaea was first collected around 1840 by Ludwig Preiss
Ludwig Preiss
Johann August Ludwig Preiss was a German-born British botanist and zoologist.Preiss was born in Herzberg am Harz, Germany. He obtained a doctorate, probably at Hamburg, then emigrated to Western Australia...

 and James Drummond
James Drummond (botanist)
James Drummond was a botanist and naturalist who was an early settler in Western Australia.-Early life:...

. For many years it was included in B. sphaerocarpa, but by 1980 it was recognised as a distinct species. In recognition of its distinctness from, yet affinity with, B. sphaerocarpa, it was for a time informally referred to as Banksia aff. Sphaerocarpa. It was eventually published by Alex George
Alex George
Alexander Segger George is a Western Australian botanist. He is the authority on the plant genera Banksia and Dryandra...

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

, based on a specimen collected by him on the Brand Highway
Brand Highway
Brand Highway is a 362 kilometre main highway linking the northern outskirts of Perth, the capital of Western Australia, to the port city of Geraldton in Western Australia's Mid West region. Together with North West Coastal Highway, it forms part of the Western Australian coastal link to the...

 about 45 km (28 mi) north of Regans Ford
Regans Ford, Western Australia
Regans Ford is a small town located in the Wheatbelt region of Western Australia, about north of Perth in the Shire of Dandaragan.-History:...

 on 14 May 1969, and labelled "A. S. George 9309". George gave it the specific name telmatiaea from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 stem
Word stem
In linguistics, a stem is a part of a word. The term is used with slightly different meanings.In one usage, a stem is a form to which affixes can be attached. Thus, in this usage, the English word friendships contains the stem friend, to which the derivational suffix -ship is attached to form a new...

 telmat-/τελματ- ("the mud of a pond"), in reference to its swampy habitat. Thus the full name for the species is Banksia telmatiaea A.S.George.

Infrageneric placement

George placed B. telmatiaea in subgenus Banksia
Banksia subg. Banksia
Banksia subg. Banksia is a valid botanic name for a subgenus of Banksia. As an autonym, it necessarily contains the type species of Banksia, B. serrata . Within this constraint, however, there have been various circumscriptions.-Banksia verae:B. subg...

 because its inflorescence is a typical Banksia flower spike, section Oncostylis
Banksia sect. Oncostylis
Banksia sect. Oncostylis is one of four sections of subgenus Banksia subg. Banksia. It contains those Banksia species with hooked pistils. All of the species in Oncostylis also exhibit a top-down sequence of flower anthesis, except for Banksia nutans which is bottom-up.Banksia sect...

 because it has hooked styles, and series Abietinae
Banksia ser. Abietinae
Banksia ser. Abietinae is avalid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

 because its inflorescence is roughly spherical. He considered its closest relative to be B. leptophylla
Banksia leptophylla
The Slender-leaved Banksia is a species of shrub in the plant genus Banksia. It occurs along the west coast of Western Australia from Gingin to Kalbarri. Before Alex George's revision of 1981, it was labelled informally as B. sphaerocarpa var. pinifolia or var...

(Slender-leaved Banksia), which differs from B. telmatiaea in having longer leaves and larger flowers; yet in his arrangement he placed it between B. scabrella
Banksia scabrella
Banksia scabrella, commonly known as the Burma Road Banksia, is a species of woody shrub in the genus Banksia. It is classified in the series Abietinae, a group of several species of shrubs with small round or oval inflorescences...

(Burma Road Banksia) and B. laricina
Banksia laricina
The Rose-Fruited Banksia is a species of shrub in the plant genus Banksia endemic to southwestern Western Australia. It derives its specific Latin name from larix or larch, which its foliage is said to resemble. The common name comes from the striking fruits which resemble wooden roses...

(Rose-fruited Banksia).

In 1996, Kevin Thiele
Kevin Thiele
Kevin R. Thiele is curator of the Western Australian Herbarium. His research interests include the systematics of the plant families Proteaceae, Rhamnaceae and Violaceae, and the conservation ecology of grassy woodland ecosystems...

 and Pauline Ladiges published the results of a cladistic
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 analysis of morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 characters of Banksia. They retained George's subgenera and many of his series, but discarded his sections. B. ser. Abietinae was found to be very nearly monophyletic
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

, and so retained. It further resolved into four subclades, so Thiele and Ladiges split it into four subseries. B. telmatiaea appeared in the third of these:

This clade became the basis of B. subser. Leptophyllae
Banksia subser. Leptophyllae
Banksia subser. Leptophyllae is avalid botanic name for a subseries of Banksia. It was published by Kevin Thiele in 1996, but discarded by Alex George in 1999.-Cladistics:...

, which Thiele defined as containing those species with "indurated and spinescent common bracts on the infructescence axes, and densely arachnose seedling stems." In accordance with their cladogram, their arrangement placed B. telmatiaea next to B. scabrella.

Thiele and Ladiges' arrangement was not accepted by George, and was largely discarded by him in his 1999 arrangement
George's taxonomic arrangement of Banksia
Alex George's taxonomic arrangement of Banksia was the first modern-day arrangement for that genus. First published in 1981 in the classic monograph The genus Banksia L.f. , it superseded the arrangement of George Bentham, which had stood for over a hundred years. It was overturned in 1996 by Kevin...

. B. ser. Abietinae was restored to George's 1981 circumscription, and all of Thiele and Ladiges' subseries were abandoned. However, B. telmatiaea was moved in the phyletic order to between B. grossa
Banksia grossa
Banksia grossa, commonly known as Coarse Banksia is a species of shrub in the plant family Proteaceae endemic to south west Western Australia. One of fourteen species of banksia with predominantly round or oval inflorescences of the series Abietinae, it was described in 1981 as a distinct species...

(Coarse Banksia) and B. leptophylla, thus better according with the affinity with B. leptophylla claimed by George in 1981.

The placement of B. telmatiaea in George's 1999 arrangement may be summarised as follows:
Banksia
Banksia
Banksia is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes and fruiting "cones" and heads. When it comes to size, banksias range from prostrate woody shrubs to trees up...

B. subg. Banksia
Banksia subg. Banksia
Banksia subg. Banksia is a valid botanic name for a subgenus of Banksia. As an autonym, it necessarily contains the type species of Banksia, B. serrata . Within this constraint, however, there have been various circumscriptions.-Banksia verae:B. subg...

B. sect. Banksia
Banksia sect. Banksia
Banksia sect. Banksia is one of four sections of Banksia subgenus Banksia. It contains those species of subgenus Banksia with straight or sometimes curved but not hooked styles. These species all have cylindrical inflorescences and usually exhibit a bottom-up sequence of flower anthesis...

 (9 series, 50 species, 9 subspecies, 3 varieties)
B. sect. Coccinea (1 species)
B. sect. Oncostylis
Banksia sect. Oncostylis
Banksia sect. Oncostylis is one of four sections of subgenus Banksia subg. Banksia. It contains those Banksia species with hooked pistils. All of the species in Oncostylis also exhibit a top-down sequence of flower anthesis, except for Banksia nutans which is bottom-up.Banksia sect...

B. ser. Spicigerae
Banksia ser. Spicigerae
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...

 (7 species, 2 subspecies, 4 varieties)
B. ser. Tricuspidae (1 species)
B. ser. Dryandroideae
Banksia ser. Dryandroideae
Banksia ser. Dryandroideae is a valid botanic name for a taxonomic series in the plant genus Banksia. First published by Carl Meissner in 1856, the name has had two circumscriptions. As presently circumscribed it is monotypic, containing only B. dryandroides.-According to Meissner:B. ser...

 (1 species)
B. ser. Abietinae
Banksia ser. Abietinae
Banksia ser. Abietinae is avalid botanic name for a series of Banksia. First published by Carl Meissner in 1856, the name has had three circumscriptions.-According to Meissner:...

B. sphaerocarpa
Banksia sphaerocarpa
Banksia sphaerocarpa, commonly known as the Fox Banksia or Round-fruit Banksia, is a species of shrub or tree in the plant genus Banksia . It is generally encountered as a 1–2 m high shrub, and is usually smaller in the north of its range...

(3 varieties)
B. micrantha
Banksia micrantha
Banksia micrantha is a species of shrub in the plant genus Banksia. A small spreading bush with pale yellow flower spikes, it occurs between Eneabba and Cervantes in South west Western Australia...

B. grossa
Banksia grossa
Banksia grossa, commonly known as Coarse Banksia is a species of shrub in the plant family Proteaceae endemic to south west Western Australia. One of fourteen species of banksia with predominantly round or oval inflorescences of the series Abietinae, it was described in 1981 as a distinct species...

B. telmatiaea
B. leptophylla
Banksia leptophylla
The Slender-leaved Banksia is a species of shrub in the plant genus Banksia. It occurs along the west coast of Western Australia from Gingin to Kalbarri. Before Alex George's revision of 1981, it was labelled informally as B. sphaerocarpa var. pinifolia or var...

(2 varieties)
B. lanata
Banksia lanata
The Coomallo Banksia is a species of shrub in the plant genus Banksia. It occurs within a range of less than 100 square kilometres between Eneabba and Mount Lesueur, Western Australia. It has roughly spherical inflorescences with flowers of cream to orange-brown colour. The leaves are linear and...

B. scabrella
Banksia scabrella
Banksia scabrella, commonly known as the Burma Road Banksia, is a species of woody shrub in the genus Banksia. It is classified in the series Abietinae, a group of several species of shrubs with small round or oval inflorescences...

B. violacea
Banksia violacea
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 high with fine narrow leaves, and is best known for its unusually coloured dark purple-violet inflorescences...

B. incana
Banksia incana
The Hoary Banksia is a species of small shrub in the plant genus Banksia. It occurs on sandplain heathland between Badgingarra and Eneabba in Western Australia, with outlying populations as far south as Perth. Seeds do not require any treatment, and take around 14 days to germinate....

B. laricina
Banksia laricina
The Rose-Fruited Banksia is a species of shrub in the plant genus Banksia endemic to southwestern Western Australia. It derives its specific Latin name from larix or larch, which its foliage is said to resemble. The common name comes from the striking fruits which resemble wooden roses...

B. pulchella
Banksia pulchella
The Teasel Banksia is a species of small shrub in the plant genus Banksia. It occurs on the south coast of Western Australia from Fitzgerald River National Park east to Israelite Bay....

B. meisneri
Banksia meisneri
The Meisner's Banksia is a species of small shrub in the plant genus Banksia. It occurs in a number of isolated populations throughout southwest Western Australia. Seeds do not require any treatment, and take 28 to 39 days to germinate.-External links:...

(2 subspecies)
B. nutans
Banksia nutans
Banksia nutans, commonly known as Nodding Banksia, is a species of shrub native to the south coast of Western Australia in the genus Banksia...

(2 varieties)
B. subg. Isostylis
Banksia subg. Isostylis
Banksia subg. Isostylis is a subgenus of Banksia. It contains three closely related species, all of which occur only in Southwest Western Australia. Members of subgenus Isostylis have dome-shaped flower heads that are superficially similar to those of B. ser...

 (3 species)


Since 1998, Austin Mast
Austin Mast
Austin R. Mast is a research botanist. Born in 1972, he obtained a Ph.D. from the University of Wisconsin–Madison in 2000. He is currently an associate professor within the Department of Biological Science at Florida State University , and has been director of FSU's since August 2003.One of his...

 has been publishing results of ongoing cladistic analyses of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 data for the subtribe Banksiinae. His analyses suggest a phylogeny that is very different to George's arrangement. With respect to B. telmatiaea, Mast's results accord closely with Thiele and Ladiges' arrangement, inferring a polytomous clade consisting of B. leptophylla, B. telmatiaea, B. scabrella and B. lanata, with B. grossa
Banksia grossa
Banksia grossa, commonly known as Coarse Banksia is a species of shrub in the plant family Proteaceae endemic to south west Western Australia. One of fourteen species of banksia with predominantly round or oval inflorescences of the series Abietinae, it was described in 1981 as a distinct species...

(Coarse Banksia) as the nearest outgroup
Outgroup
In cladistics or phylogenetics, an outgroup is a group of organisms that serves as a reference group for determination of the evolutionary relationship among three or more monophyletic groups of organisms....

:

Early in 2007, Mast and Thiele initiated a rearrangement of Banksia by merging Dryandra
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 extensive molecular and morphological evidence that Banksia was paraphyletic with...

into it, and publishing B. subg. Spathulatae
Banksia subg. Spathulatae
Banksia subg. Spathulatae is a valid botanic name for a subgenus of Banksia. It was published in 2007 by Austin Mast and Kevin Thiele, and defined as containing all those Banksia species having spathulate cotyledons...

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

s. 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, then B. telmatiaea is placed in B. subg. Spathulatae.

Distribution and habitat

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

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

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

 biogeographic
Biogeography
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...

 regions, inland from the coast but never east of the Darling Scarp
Darling Scarp
The Darling Scarp, also referred to as the Darling Range or Darling Ranges, is a low escarpment running north-south to the east of the Swan Coastal Plain and Perth, Western Australia...

. It occurs from Hill River
Hill River (Western Australia)
Hill River is a river and town in the Wheatbelt region of Western Australia. At the 2006 census, Hill River had a population of 134.The headwaters of the river below Dinner Hill approximately East of Badgingarra then flows in a westerly direction crossing the Brand Highway just North of...

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

 in the north, to Serpentine
Serpentine, Western Australia
Serpentine is a town located south-southeast of Perth, the capital of Western Australia, and 7 km south of Mundijong. Serpentine is located on the South Western Railway between Perth and Bunbury, and was one of the original stations when the line was opened in 1893...

 in the south. Most populations occur north of Moore River
Moore River
Moore River in Western Australia can refer to a number of places:*Moore River - the river itself*Moore River National Park - national park that the river runs through...

 or south of Cannington
Cannington, Western Australia
Cannington is a southern suburb of Perth, Western Australia. Its Local Government Area is the City of Canning.-History:Cannington's name derives from the Canning River, which forms part of the southwestern boundary of the suburb...

, there being only a few scattered populations in between.

The species favours lowland areas that are seasonally wet but never inundated, such as the margins of swamp
Swamp
A swamp is a wetland with some flooding of large areas of land by shallow bodies of water. A swamp generally has a large number of hammocks, or dry-land protrusions, covered by aquatic vegetation, or vegetation that tolerates periodical inundation. The two main types of swamp are "true" or swamp...

s and marshes. For example, in the Yule Brook Botany Reserve
Yule Brook Botany Reserve
Yule Brook Botany Reserve, also known as Yule Brook Reserve and Cannington Swamps, is a 34.6 hectare parcel of land in the Perth, Western Australia suburb of Kenwick. It is owned by the University of Western Australia, and used by them for botanical research and teaching.It was purchased by the...

, where parallel sand ridge
Ridge
A ridge is a geological feature consisting of a chain of mountains or hills that form a continuous elevated crest for some distance. Ridges are usually termed hills or mountains as well, depending on size. There are several main types of ridges:...

s cross a clay flat, B. telmatiaea occurs neither in the lowest parts of the flat, where seasonal inundation occurs; nor on the tops of the ridges, where the drainage is good; but it is one of the most abundant plants of intermediate habitats, on ridge slopes and in higher areas of the clay flat.

Favoured soils are deep grey sandy loams or shallower sand overlying claypan
Claypan
In geology, a claypan is a dense, compact, slowly permeable layer in the subsoil having a much higher clay content than the overlying material, from which it is separated by a sharply defined boundary. Claypans are usually hard when dry, and plastic and sticky when wet. They limit or slow the...

. Associated vegetation is typically scrubland or shrubland
Shrubland
Shrubland, scrubland, scrub or brush 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...

, although moisture-loving trees such as B. littoralis
Banksia littoralis
Banksia littoralis, commonly known as the Swamp Banksia, Swamp Oak, Pungura and the Western Swamp Banksia, is a tree in the plant genus Banksia. It is found in south west Western Australia from the south eastern metropolitan area of Perth to the Stirling Range and Albany...

(Swamp Banksia) or Melaleuca preissiana
Melaleuca preissiana
Melaleuca preissiana, commonly known as Stout Paperbark, Modong or Moonah, is a tree that grows in coastal areas of southwest Australia. It grows up to 15 metres tall, occurring chiefly in areas that are seasonally wet. It has papery bark, and pointed leaves from 1 to 1½ centimetres long and 1 to 2...

(Moonah) may also be present, sometimes in sufficient numbers to form a low open woodland
Woodland
Ecologically, a woodland is a low-density forest forming open habitats with plenty of sunlight and limited shade. Woodlands may support an understory of shrubs and herbaceous plants including grasses. Woodland may form a transition to shrubland under drier conditions or during early stages of...

.

Ecology

Like most other Proteaceae
Proteaceae
Proteaceae is a family of flowering plants distributed in the Southern Hemisphere. The family comprises about 80 genera with about 1600 species. Together with the Platanaceae and Nelumbonaceae they make up the order Proteales. Well known genera include Protea, Banksia, Embothrium, Grevillea,...

, B. telmatiaea has proteoid root
Proteoid root
Proteoid roots, also known as cluster roots, are plant roots that form clusters of closely spaced short lateral rootlets. They may form a two to five centimetre thick mat just beneath the leaf litter. They enhance nutrient uptake, possibly by chemically modifying the soil environment to improve...

s, roots with dense clusters of short lateral rootlets that form a mat in the soil just below the leaf litter. These roots are particularly efficient at absorbing nutrients from nutrient-poor soils, such as the phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

-deficient native soils of Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

.

Unlike many Banksia species, B. telmatiaea lacks a lignotuber
Lignotuber
A lignotuber is a starchy swelling of the root crown possessed by some plants as a protection against destruction of the plant stem by fire. The crown contains buds from which new stems may sprout, and a sufficient store of nutrients to support a period of growth in the absence of...

, so plants are killed by bushfire. However, it is adapted to release its aerial seed bank following a bushfire, and so regenerates rapidly. This behaviour, known as serotiny
Serotiny
Serotiny is an ecological adaptation exhibited by some seed plants, in which seed release occurs in response to an environmental trigger, rather than spontaneously at seed maturation. The most common and best studied trigger is fire, and the term serotiny is often used to refer to this specific case...

, makes B. telmatiaea dependent upon a suitable fire regime for successful regeneration; indeed, excessive fire frequency may be one reason why B. telmatiaea does not occur further south, despite suitable habitat throughout southwest Australia
Southwest Australia
Southwest Australia is a biodiversity hotspot that includes the Mediterranean forests, woodlands, and scrub ecoregions of Western Australia. The region has a wet-winter, dry-summer Mediterranean climate, one of five such regions in the world...

. Unlike most serotinous Banksia species, the seeds of B. telmatiaea are not released immediately after the passage of a bushfire. The follicles open straight away, but at first the seeds are blocked from falling out by the winged seed separator
Seed separator
A seed separator is a structure found in the follicles of some Proteaceae. These follicles typically contain two seeds, with a seed separator between them...

. If moistened, these wings close up, and as they dry they open out again, levering the seeds out of position, making it possible for them to fall. This adaptation ensures that seeds are released only after the first rains following a bushfire.

Four species of bird have been observed visiting the flowers of B. telmatiaea: Anthochaera carunculata (Red Wattlebird), Zosterops lateralis (Silvereye), Phylidonyris novaehollandiae (New Holland Honeyeater) and the Lichmera indistincta (Brown Honeyeater). The introduced Apis mellifera (European Honey Bee) is also commonly observed, and visits by ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s and Hylaeus
Hylaeus (genus)
Hylaeus is a large and diverse cosmopolitan genus within the bee family Colletidae, consisting of generally small, black and yellow/white wasp-like species...

plasterer bees have been recorded. Visits by nectarivorous
Nectarivore
In zoology, nectarivore is an animal which eats the sugar-rich nectar produced by flowering plants. Most nectarivores are insects or birds, but there are also nectarivorous mammals, notably several species of bats in the Southwestern United States and Mexico, as well as the Australian Honey Possum...

 mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s have not been directly observed, but their involvement in pollination is certain, as their scats
Feces
Feces, faeces, or fæces is a waste product from an animal's digestive tract expelled through the anus or cloaca during defecation.-Etymology:...

 have often been found on inflorescences, and studies of other Banksia species have consistently demonstrated their involvement. Moreover, a number of characteristics of the B. telmatiaea spike are purported to be adaptations to pollination by nocturnal mammals: the strong, musky odour, the occurrence of inflorescences hidden within the foliage close to the ground, the large amounts of nectar produced, and the pattern of nectar production, which peaks at dawn and dusk. This last adaptation is thought to favour visits by bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s and mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s, which feed in the morning and evening respectively, as opposed to insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, which are most active during the day.

Reproductive success is strongly affected by insects that infest the flower spikes and fruiting structures. Infestation of the flower spikes is not as severe as in other Banksia species: one study found less than 10% of B. telmatiaea inflorescences to be infested, compared to over 50% for B. attenuata
Banksia attenuata
Banksia attenuata, commonly known as the candlestick banksia or slender banksia, is a species of plant in the proteaceae family. Commonly a tree, it reaches 10 m high, but is often a shrub in dryer areas 0.4 to 2 m high...

(Candlestick Banksia), B. littoralis and B. menziesii
Banksia menziesii
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 tall, or a lower spreading 1–3 m shrub in the more northern parts of its range. The serrated leaves are dull green with new growth a paler grey...

(Menzies' Banksia), and over 90% for B. grandis
Banksia grandis
Banksia grandis, commonly known as Bull Banksia, Giant Banksia or Mangite, is a common and distinctive tree in South West Western Australia....

(Bull Banksia). Also, whereas other species were attacked by a range of insects, the inflorescence of B. telmatiaea was attacked only by the tortrix
Tortrix
Tortrix is a genus of moths belonging to the Tortricidae family.-Species:*Tortrix sinapina *Tortrix viridana Tortrix? destructus Tortrix? florissantana...

 moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

 Arotrophora arcuatalis
Arotrophora arcuatalis
Arotrophora arcuatalis, commonly known as Banksia Boring Moth or rarely Banksia Moth, is a species of Australian tortrid moth best known as a pest of Banksia....

(Banksia Boring Moth), which burrows into the woody axis, rendering the spike barren. On the other hand, the same study observed heavy infestation of fruiting structures, with over 90% of spikes with follicles found to contain at least one larva of an unidentified species of Xylorycta
Xylorycta
Xylorycta is a genus of Australian oecophorid moth. Xylorycta species are strongly associated with the plant family Proteaceae, being found on Hakea, Lambertia, Grevillea, Leptospermum, Macadamia, Oreocallis, Persoonia and Telopea...

. These larvae burrow from follicle to follicle to eat the seed, resulting in 100% seed loss for infested spikes.

B. telmatiaea is one of five Banksia species, all closely related to B. sphaerocarpa, that have highly unusual flower nectar. Whereas other Banksia species produce nectar that is clear and watery, the nectar of these species is pale yellow initially, but gradually becomes darker and thicker, changing to a thick, olive-green mucilage within one to two days of secretion. In the case of B. telmatiaea, it eventually becomes "an almost black, gelatinous lump adhering to the base of the flowers". This unusual nectar was first noted in 1980 by Byron Lamont
Byron Lamont
Professor Byron Barnard Lamont is a Western Australian botanist. He is currently a senior researcher within the Department of Environmental Biology of Curtin University of Technology...

, who attributed it to the cyanobacteria that he observed feeding off the nectar sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

s. Noting that many of these cyanobacteria had heterocyst
Heterocyst
Heterocysts are specialized nitrogen-fixing cells formed by some filamentous cyanobacteria, such as Nostoc punctiforme, Cylindrospermum stagnale and Anabaena sphaerica, during nitrogen starvation. They fix nitrogen from dinitrogen in the air using the enzyme nitrogenase, in order to provide the...

s, he speculated that they aid the plant by fixing atmospheric nitrogen
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

, which is then washed off the flower heads by rain, and absorbed by the proteoid root mat. This purported symbiosis
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 was investigated in 1985, but no evidence of nitrogen fixing was found. Further investigations in 1996 suggested that the discolouration is not caused by cyanobacteria or other microorganisms in the nectar, but is rather "a chemical phenomenon of plant origin". As of February 2007, the cause was still unknown. Chemical analysis of B. telmatiaea nectar has shown it to have normal nectar sugar compositions, albeit sucrose
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...

-dominant.

Conservation

B. telmatiaea is a fairly secure species, as most populations are of more than 100 plants, and 26% of known plants are in conservation reserves. Its proximity to Perth
Perth, Western Australia
Perth is the capital and largest city of the Australian state of Western Australia and the fourth most populous city in Australia. The Perth metropolitan area has an estimated population of almost 1,700,000....

 suggests that land clearing for urban development could pose a threat, and in 1988 The Banksia Atlas
The Banksia Atlas
The Banksia Atlas is an atlas that documents the ranges, habitats and growth forms of various species and other subgeneric taxa of Banksia, an iconic Australian wildflower genus...

recommended that "the species should continue to be monitored since land clearing could change the situation greatly, particularly amongst its northern populations." It is also known to be susceptible to dieback caused by the introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 plant pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

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

, a soil-borne water mould
Water mould
Oömycota or oömycetes form a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms . They are filamentous, microscopic, absorptive organisms that reproduce both sexually and asexually...

 that causes root rot; in fact it is so reliably susceptible that it used as an indicator species for the presence of the disease. An assessment of the potential impact of climate change
Climate change
Climate change is a significant and lasting change in the statistical distribution of weather patterns over periods ranging from decades to millions of years. It may be a change in average weather conditions or the distribution of events around that average...

 on this species found that severe change is likely to lead to extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...

; but under less severe change scenarios the distribution may actually grow, depending on how effectively it can migrate into newly habitable areas.

In 1987, George applied the Rare or Threatened Australian Plants (ROTAP
ROTAP
Rare or Threatened Australian Plants, usually abbreviated to ROTAP, is a list of rare or threatened Australian plant taxa. Developed and maintained by the CSIRO, the most recent edition lists 5031 taxa. The list uses a binary coding system based on the IUCN Red List categories for "Presumed...

) criteria to the species, determining it to have a conservation status of "3R": a rare species found only in small populations, but not considered endangered or vulnerable. Western Australia's Department of Environment and Conservation do not consider it to be rare, however, and have not included it on their Declared Rare and Priority Flora List.

Cultivation

B. telmatiaea is rarely cultivated. It grows fairly quickly, but tends to become untidy as it ages. The flower spikes, though attractive, occur within the bush where they are usually obscured by foliage. In its natural habitat it flowers prolifically over several months, but according to George it may be reluctant to flower in cultivation. It tolerates light pruning not below the green foliage. George recommends a sunny position in poorly drained soil, preferably with moisture in winter. Seeds do not require any treatment, and take around 14 days to germinate.

External links

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