Gevuininae
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The Grevilleoideae are a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
of the plant family
Proteaceae The Proteaceae form a family of flowering plants predominantly distributed in the Southern Hemisphere. The family comprises 83 genera with about 1,660 known species. Together with the Platanaceae and Nelumbonaceae, they make up the order Pro ...
. Mainly restricted to the Southern Hemisphere, it contains around 46
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and about 950
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. Genera include ''
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. ''Banksias'' range i ...
'', ''
Grevillea ''Grevillea'', commonly known as spider flowers, is a genus of about 360 species of evergreen flowering plants in the family Proteaceae. Plants in the genus ''Grevillea'' are shrubs, rarely trees, with the leaves arranged alternately along the b ...
'', and ''
Macadamia ''Macadamia'' is a genus of four species of trees in the flowering plant family Proteaceae. They are indigenous to Australia, native to northeastern New South Wales and central and southeastern Queensland specifically. Two species of the genus ...
''.


Description

The Grevilleoideae grow as
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s,
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
s, or
subshrub A subshrub (Latin ''suffrutex'') or dwarf shrub is a short shrub, and is a woody plant. Prostrate shrub is a related term. "Subshrub" is often used interchangeably with "bush".Jackson, Benjamin, Daydon; A Glossary of Botanic Terms with their Der ...
s. They are highly variable, making a simple, diagnostic identification key for the subfamily essentially impossible to provide. One common and fairly diagnostic characteristic is the occurrence of flowers in pairs that share a common
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
. However, a few Grevilleoideae taxa do not have this property, having solitary flowers or
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. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s of unpaired flowers. In most taxa, the flowers occur in densely packed heads or spikes, and the
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
is a follicle.


Distribution and habitat

Grevilleoideae are mainly a Southern Hemisphere family. The main centre of diversity is
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, with around 700 of 950 species occurring there, and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
also contains taxa. However, the Grevilleoideae are barely present in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
; almost all of the Proteaceae taxa there belong to the subfamily
Proteoideae Proteoideae is one of the five subfamilies of the plant family Proteaceae. The greatest diversity is in Africa, but there are also many species in Australia; a few species occur in South America, New Caledonia, and elsewhere. Taxonomy Proteoidea ...
. The ''Brabejum'' tree of
Cape Town Cape Town ( af, Kaapstad; , xh, iKapa) is one of South Africa's three capital cities, serving as the seat of the Parliament of South Africa. It is the legislative capital of the country, the oldest city in the country, and the second largest ...
is the exception, and the only grevilleoid in Africa.


Taxonomy

The framework for classification of the Proteaceae was laid by L.A.S. Johnson and Barbara Briggs in their 1975 monograph " On the Proteaceae: the evolution and classification of a southern family". Their classification has been refined somewhat over the ensuing three decades, most notably by
Peter H. Weston Peter may refer to: People * List of people named Peter, a list of people and fictional characters with the given name * Peter (given name) ** Saint Peter (died 60s), apostle of Jesus, leader of the early Christian Church * Peter (surname), a su ...
and Nigel Barker in 2006. The Grevilleoideae are now considered one of five subfamilies of the Proteaceae. The placement and circumscription of the Grevilleoideae in four
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...
, according to Weston and Barker can be summarised as: * Tribe
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
::::'' Sphalmium'' — '' Carnarvonia''


Roupaleae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...

Authority:
Meisn. Carl Daniel Friedrich Meissner (1 November 1800 – 2 May 1874) was a Swiss botanist. Biography Born in Bern, Switzerland on 1 November 1800, he was christened Meisner but later changed the spelling of his name to Meissner. For most of his 40 ...
:::
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
::::''
Megahertzia ''Megahertzia'' is a monotypic genus of flowering plants in the family Proteaceae. The sole species, ''Megahertzia amplexicaulis'', is endemic to Queensland. It is found only in part of the Wet Tropics World Heritage Area between the Daintree Ri ...
'' — ''
Knightia ''Knightia'' is an extinct genus of clupeid bony fish that lived in the freshwater lakes and rivers of North America and Asia during the Eocene epoch. The genus was erected by David Starr Jordan in 1907, in honor of the late University of Wyom ...
'' — ''
Eucarpha ''Eucarpha'' is a genus of flowering plant of the family Proteaceae, endemic to New Caledonia. Two species are recognised. Up to 1975, these were classified within the genus ''Knightia (plant), Knightia'' until Lawrence Alexander Sidney Johnson, ...
'' — ''
Triunia ''Triunia'' is a genus of medium to tall shrubs or small trees found as understorey plants in rainforests of eastern Australia. Members of the plant family Proteaceae, they are notable for their poisonous fleshy fruits or drupes. Only one species ...
'' :::Subtribe
Roupalinae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Roupala ''Roupala'' is a Neotropical genus of woody shrubs and trees in the plant family Proteaceae. Its 34 species are generally found in forests from sea level to 4000 m altitude from Mexico to Argentina. Taxonomy and naming The genus was described b ...
'' — ''
Neorites ''Neorites'' is a monotypic genus of plants in the family Proteaceae. The sole species ''Neorites kevedianus'', commonly called fishtail oak or fishtail silky oak, is a tall tree endemic to the wet tropics rainforests of north eastern Queenslan ...
'' — ''
Orites ''Orites'' is a genus of 9 known species, 7 endemic to Australia (4 of which occur in Tasmania) and 2 in South America; 1 in the Chilean Andes and 1 in Bolivia. Species This listing was sourced from the ''Australian Plant Name Index'' and ot ...
'' :::Subtribe Lambertiinae ( Venk.Rao) L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Lambertia ''Lambertia'' is a genus of flowering plants, belonging to the family Proteaceae. It is endemic to Australia. The genus was named in 1798 by Sir James Edward Smith in honour of English botanist Aylmer Bourke Lambert. The ''Lambertias'' are scl ...
'' — ''
Xylomelum ''Xylomelum'' is a genus of six species of flowering plants, often commonly known as woody pears, in the family Proteaceae and are endemic to Australia. Plants in this genus are tall shrubs or small trees with leaves arranged in opposite pairs, ...
'' :::Subtribe
Heliciinae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Helicia ''Helicia'' is a genus of 110 species of trees and shrubs, constituting part of the plant family Proteaceae. They grow naturally in rainforests throughout tropical South and Southeast Asia, including India, Sri Lanka, Indochina, Peninsular Mala ...
'' — '' Hollandaea'' :::Subtribe
Floydiinae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Darlingia ''Darlingia'' is a small genus of two species of rainforest tree from Northern Queensland. It was described by Ferdinand von Mueller Baron Sir Ferdinand Jacob Heinrich von Mueller, (german: Müller; 30 June 1825 – 10 October 1896) was a ...
'' — ''
Floydia ''Floydia'' is a monotypic species of tree in the family Proteaceae native to Australia. It is a somewhat rare tree found only growing in the rainforests of southeastern Queensland and northern New South Wales. The sole species is ''Floydia prae ...
''


Banksieae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genus, genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Descriptio ...

Authority: Rchb. :::fossil form genera ::::'' Banksieaeidites''  — ''
Banksieaeformis ''Banksieaeformis'' is a genus that encompasses plant species only known from fossil leaves that can be attributed to the Proteaceae tribe Banksieae, but cannot be attributed to an extant (living) genus. Unlike those classified in the related gen ...
''  — ''
Banksieaephyllum ''Banksieaephyllum'' is a plant genus that encompasses organically preserved fossil leaves that can be attributed to the Proteaceae tribe Banksieae, but cannot be attributed to a genus. Before 1950, many fossil leaves were attributed to the gen ...
'' :::Subtribe
Musgraveinae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genus, genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Descriptio ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Musgravea ''Musgravea'' is a genus of rainforest tree from north-eastern Queensland. It was published in 1890 by Ferdinand von Mueller, and named in honour of Sir Anthony Musgrave, Governor of Queensland from 1883 to 1888. Together with its closest rel ...
'' — ''
Austromuellera ''Austromuellera'' is a genus of only two known species of medium-sized trees, constituting part of the plant family Proteaceae. They are both endemic to three restricted areas of the wet tropics rainforests of north-eastern Queensland, Austra ...
'' :::Subtribe
Banksiinae ''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 ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
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. ''Banksias'' range i ...
''


Embothrieae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...

Authority: Rchb. :::Subtribe Lomatiinae L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Lomatia ''Lomatia'' is a genus of 12 species of evergreen flowering plants in the protea family Proteaceae. Within the family, they have been placed, alone, in their own subtribe, Lomatiinae according to Johnson & Briggs 1975 classification of the fam ...
'' :::Subtribe
Embothriinae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description T ...
Endl. Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Pressburg, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian Empire, Austrian botanist, numismatist and Sinologist. He was a director of the Botan ...
::::''
Embothrium ''Embothrium'' is a genus of two to eight species (depending on taxonomic interpretation) in the plant family Proteaceae, native to southern South America, in Chile and adjacent western Argentina; the genus occurs as far south as Tierra del Fuego ...
'' — ''
Oreocallis ''Oreocallis'' is a South American plant genus in the family Proteaceae. There is only one species, ''Oreocallis grandiflora'', which is native to mountainous regions in Peru and Ecuador. Previously, the genus was considered to have several spec ...
'' — ''
Alloxylon ''Alloxylon'' is a genus of four species in the family Proteaceae of mainly small to medium-sized trees. They are native to the eastern coast of Australia, with one species, '' A. brachycarpum'' found in New Guinea and the Aru Islands. The genus ...
'' — '' Telopea'' :::Subtribe Stenocarpinae L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Stenocarpus ''Stenocarpus'' is a genus of about 22 species of flowering plants in the family Proteaceae. They are trees or shrubs with variably-shaped leaves, zygomorphic, bisexual flowers, the floral tube opening on the lower side before separating i ...
'' — '' Strangea'' :::Subtribe Hakeinae
Endl. Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Pressburg, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian Empire, Austrian botanist, numismatist and Sinologist. He was a director of the Botan ...
::::''
Opisthiolepis ''Opisthiolepis'' is a genus of a sole described species of large trees, constituting part of the plant family Proteaceae. The species ''Opisthiolepis heterophylla'' most commonly has the names of blush silky oak, pink silky oak, brown silky oa ...
'' — ''
Buckinghamia ''Buckinghamia'' is a genus of only two known species of trees, belonging to the plant family Proteaceae. They are endemic to the rainforests of the wet tropics region of north eastern Queensland, Australia. The ivory curl flower, ''B. celsissi ...
'' — ''
Hakea ''Hakea'' ( ) is a genus of about 150 species of plants in the Family ''Proteaceae'', endemic to Australia. They are shrubs or small trees with leaves that are sometimes flat, otherwise circular in cross section in which case they are sometimes d ...
'' — ''
Grevillea ''Grevillea'', commonly known as spider flowers, is a genus of about 360 species of evergreen flowering plants in the family Proteaceae. Plants in the genus ''Grevillea'' are shrubs, rarely trees, with the leaves arranged alternately along the b ...
'' — '' Finschia''


Macadamieae

Authority: Venk.Rao :::Subtribe Macadamiinae L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Macadamia ''Macadamia'' is a genus of four species of trees in the flowering plant family Proteaceae. They are indigenous to Australia, native to northeastern New South Wales and central and southeastern Queensland specifically. Two species of the genus ...
'' — ''
Panopsis ''Panopsis'' is a genus of trees in the family Proteaceae. The species, which occur in Central and South America, include: Newer species that can be considered part of the ''Panopsis'' genus has been discovered recently, called ''Panopsis magnif ...
'' — ''
Brabejum ''Brabejum'' is a genus of a single species of large evergreen tree, ''Brabejum stellatifolium'' in the family Proteaceae, commonly called wild almond, bitter almond or ghoeboontjie. It is restricted in the wild to South Africa's Western Cape pro ...
'' :::Subtribe Malagasiinae P.H.Weston & N.P.Barker ::::'' Malagasia'' — ''
Catalepidia ''Catalepidia'' is a genus of a sole described species of medium-sized trees, constituting part of the plant family Proteaceae. The species ''Catalepidia heyana'' grows naturally only in a restricted mountain region ( endemic) of the wet trop ...
'' :::Subtribe Virotiinae P.H.Weston & N.P.Barker ::::'' Virotia'' — ''
Athertonia ''Athertonia'' is a monotypic genus of plants in the family Proteaceae. The sole described species is ''Athertonia diversifolia'', commonly known as Atherton oak, athertonia, creamy silky oak or white oak. It is endemic to a small part of the ...
'' — ''
Heliciopsis ''Heliciopsis'' is a genus of about thirteen species of trees, constituting part of the flowering plant family Proteaceae. They grow naturally in Burma, Indo-China, SE. China, Thailand, Peninsular Malaysia, Borneo, Sumatra, Java (Indonesia) and ...
'' :::Subtribe
Gevuininae The Grevilleoideae are a subfamily of the plant family Proteaceae. Mainly restricted to the Southern Hemisphere, it contains around 46 genera and about 950 species. Genera include ''Banksia'', ''Grevillea'', and ''Macadamia''. Description The ...
L.A.S.Johnson &
B.G.Briggs Barbara Gillian Briggs (born 1934) is one of the foremost Australian botanists. The '' IK'' lists 205 names of plants which have been published or co-published by her. She was one of the botanists in the Angiosperm Phylogeny Group, of th1998APG ...
::::''
Cardwellia ''Cardwellia'' is a genus of a sole described species of large trees in the plant family Proteaceae. The species ''Cardwellia sublimis'' (northern silky oak) is endemic to the rainforests of the wet tropics region of northeastern Queensland, A ...
'' — ''
Sleumerodendron ''Sleumerodendron'' is a monotypic genus of plant in the family Proteaceae. The sole species is ''Sleumerodendron austrocaledonicum''. This species is endemic to New Caledonia. It is closely related to ''Turrillia'' (Vanuatu, Fiji) and ''Kermade ...
'' — ''
Euplassa ''Euplassa'' is a genus of flowering plants in the protea family. It is native to tropical South America, including Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, and Venezuela."''Euplassa'' Salisb. ex Knight". Plants ...
'' — ''
Gevuina ''Gevuina avellana'' (Chilean hazelnut ( in Spanish), or ''Gevuina hazelnut''), is an evergreen tree, up to 20 meters (65 feet) tall. It is the only species currently classified in the genus ''Gevuina''. It is native to southern Chile and adjace ...
'' — ''
Bleasdalea ''Bleasdalea'' is a genus of flowering plants in the family Proteaceae. Taxonomy Molecular and morphological analysis shows this genus is most closely related to the genus '' Hicksbeachia'', ancestors of the two genera having diverged around 15 ...
'' — ''
Hicksbeachia ''Hicksbeachia'' is a genus of two species of trees in the family Proteaceae. They are native to rainforests of northern New South Wales and southeastern Queensland. They are commonly known as red bopple nut or beef nut due to the bright red col ...
'' — ''
Kermadecia ''Kermadecia'' is a genus of flowering plants in the family Proteaceae. The genus comprises five species, all endemic to New Caledonia. Its closest relatives are ''Sleumerodendron'' (New Caledonia) and ''Turrillia'' (Fiji, Vanuatu),Mast, A. R., ...
'' — ''
Turrillia '' Turrillia '' is a genus of plants in the family Proteaceae, native to Oceania. The genus was named by American botanist Albert Charles Smith in 1985, in ''Flora Vitiensis Nova'' 3, containing five species. Only three, from Fiji and Vanuatu, a ...
''


Uses

The nursery industry cultivates many Grevilleoideae species as barrier plants and for their prominent and distinctive flowers and foliage. Some species are of importance to the cut-flower industry, especially some ''
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. ''Banksias'' range i ...
'' and ''
Dryandra ''Banksia'' ser. ''Dryandra'' is a series of 94 species of shrub to small tree in the plant genus ''Banksia''. It was considered a separate genus named ''Dryandra'' until early 2007, when it was merged into ''Banksia'' on the basis of extensiv ...
'' species. Two species of the genus ''
Macadamia ''Macadamia'' is a genus of four species of trees in the flowering plant family Proteaceae. They are indigenous to Australia, native to northeastern New South Wales and central and southeastern Queensland specifically. Two species of the genus ...
'' and the Chilean species ''
Gevuina avellana ''Gevuina avellana'' (Chilean hazelnut ( in Spanish language, Spanish), or ''Gevuina hazelnut''), is an evergreen tree, up to 20 meters (65 feet) tall. It is the only species currently classified in the genus ''Gevuina''. It is native to souther ...
'' (Chilean hazel) are grown commercially for edible nuts. Chilean hazel has an acceptable frost tolerance.


References


External links

{{Taxonbar, from=Q1934685 Proteaceae Eudicot subfamilies