Difference between revisions of "Proteaceae"

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#redirect [[:Category:Proteaceae]]
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{{Automatic taxobox
 
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|name = Proteaceae
 
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|image = Protea cynaroides 3.jpg
 
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|image_caption = [[Inflorescence]] of ''[[Protea cynaroides]]''
 
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|taxon = Proteaceae
 
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|authority = [[Antoine Laurent de Jussieu|Juss.]]<ref name=APGIII2009>{{Cite journal |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |url=http://onlinelibrary.wiley.com/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/pdf | format= PDF |accessdate=2013-07-06 |doi=10.1111/j.1095-8339.2009.00996.x }}</ref>
 
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|subdivision_ranks = Genera
 
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|subdivision = About 80, see text
 
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|}}
 
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The '''Proteaceae''' {{IPAc-en|ˌ|p|r|oʊ|t|i|'|eɪ|s|iː}} are a [[family (biology)|family]] of [[flowering plants]] predominantly distributed in the [[Southern Hemisphere]]. The family comprises 83 [[genus|genera]] with about 1,660 known [[species]].<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W.  |lastauthoramp=yes | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press }}</ref> Together with the [[Platanaceae]] and [[Nelumbonaceae]], they make up the order [[Proteales]]. Well-known genera include ''[[Protea]]'', ''[[Banksia]]'', ''[[Embothrium]]'', ''[[Grevillea]]'', ''[[Hakea]]'', ''[[Dryandra]]'', and ''[[Macadamia]]''. Species such as the New South Wales waratah (''[[Telopea speciosissima]]''), king protea (''[[Protea cynaroides]]''), and various species of ''Banksia'', ''Grevillea'', and ''Leucadendron'' are popular cut flowers, while the nuts of ''[[Macadamia integrifolia]]'' are widely grown commercially and consumed. [[Australia]] and [[South Africa]] have the greatest concentrations of diversity.
 
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==Etymology==
 
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The name Proteaceae by [[Antoine Laurent de Jussieu]] in 1789 was based on the genus ''Protea'', which in 1767 [[Carl Linnaeus]] derived from the name of the Greek god [[Proteus]], a deity that was able to change between many forms. This is an appropriate image, seeing as the family is known for its astonishing variety and diversity of flowers and leaves.<!-- The original reference publication sources for this section as links to be fully cited here, in due course: → http://biodiversitylibrary.org/page/42945242 → http://biodiversitylibrary.org/page/7439108 -->
 
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== Description ==
 
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[[File:Rhopala heterophylla Pohl90.png|thumb|right|''Rhopala heterophylla'']]
 
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The genera of Proteaceae are highly varied, with ''[[Banksia]]'' in particular providing a striking example of [[adaptive radiation]] in plants.<ref name="Mast 2002">{{cite journal |author1=Mast, A. R.  |author2=Givnish, T. J.  |lastauthoramp=yes | year = 2002 | title = Historical Biogeography and the Origin of Stomatal Distributions in Banksia & Dryandra (Proteaceae) Based on Their cpDNA Phylogeny | journal = American Journal of Botany | volume = 89 | issue = 8 | pages = 1311–1323 | doi = 10.3732/ajb.89.8.1311 | pmid=21665734}}</ref> This variability makes it impossible to provide a simple, diagnostic identification key for the family, although individual genera may be easily identified.
 
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* Proteaceae are generally [[tree]]s, rarely of more than 40 m in height, and are usually of medium height or low or perennial [[shrub]]s, except for some ''[[Stirlingia]]'' species that are [[herb]]s. They are facultatively deciduous (''[[Embothrium coccineum]]''), rarely acaulescent, the cauline portion of the collar is often thickened ([[lignotuber]]). [[Indumentum]] of three-celled hairs, sometimes glandular, rarely absent, the apical cell is usually elongated, acute, sometimes equally or unequally bifid.
 
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* [[Leaf|Leaves]] rarely aromatic, usually alternate, and in a spiral, rarely opposed, or verticilate; coriaceous, rarely fleshy or spinescent, simple or compound (imparipinate, imparibipinate or rarely palmate or digitate with pinnatisect segments), entire edge to (3-)pinnatisect (giving a fern-like aspect); rarely divided dichotomously, often remotely toothed, crenate or serrated, seated or stalked; the [[Petiole (botany)|petiole]] frequently with a swollen base but rarely sheathed (sometimes in ''[[Synaphea]]''), without [[stipule]]s; pinnate sometimes palmate or parallel [[Leaf#venation|venation]], brochidodromous or reduced to a single prominent vane, [[vernation]] normally conduplicate; anisophylly often occurs during the different growth periods; leaf blade dorsiventral, isobilateral or centred; [[mesophyll tissue]] usually with sclerenchymatous [[idioblast]]s, rare secretory cavities. Brachy-paracytic [[stoma]]ta (laterocytic in ''[[Bellendena]]'').
 
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[[Plant stem]]s with two types of radii, wide and multi-serrated or narrow and uni-serrated, [[phloem]] stratified or not, trilacunar nodes with three [[Leaf gap|leaf trace]]s (rarely unilacunar with one trace), [[ground tissue|sclereids]] frequent; [[bark]] with [[lenticel]]s frequently horizontally enlarged, [[cork cambium]] present, usually superficial. [[Root]]s lateral and short, often grouped in bundles (''proteoid roots'') with very dense root hairs, rarely with [[mycorrhiza]].
 
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* [[Plant]]s usually hermaphroditic, more rarely monoecious, dioecious or andromonoecious.
 
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* [[Inflorescence]]s very variable, simple or compound, axillary or terminal, lateral flowers solitary or in pairs, rarely with a terminal flower, [[Raceme|racemiform]], [[Panicle|paniculate]] or condensed, usually with [[bract]]s, sometimes converted into leaves or squamiform, forming a type of cone, or with bright colours, forming an involucre or pseudanthium, the peduncles and [[Pedicel (botany)|pedicel]]s sometimes contracted, compacted with the [[rachis]], in some cases the congested inflorescences form super inflorescences (some ''[[Alloxylon]]''); very rarely the flowers are solitary and axillary near the end of branches; in species with lignotubers the flowers sometimes grow from these and pass through the soil (geophytes).
 
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* [[Flower]]s are usually perfect, actinomorphic, or zygomorphic, hypogynous, frequently large and showy. Flat or oblique, sometimes forming a gynophore. Hypogynous disk present and extrastaminal or absent. [[Perianth]] of (3-)4(-8) [[tepal]]s (sometimes interpreted as a dimerous and dichlamydeous perianth), in 1(-2) valvate whorls, sometimes elongated in a basal sack, free or fused in different ways (all fused or even one free and three basally to completely fused), or even connivent by marginally interdigitate papillae forming a tube or a bilabiate structure, zygomorphic, sometimes opening laterally in a variety of ways. Haplostemonous androecium, usually isostemonous, opposititepalous of (3-)4(-5) [[stamen]]s, all fertile or some converted into [[staminode]]s, usually filamentous, filaments partially or totally fused to the tepals, rarely free, basifixed [[anther]]s adnate, ditheous, tetrasporangiate, sometimes unilocular and bisporagiate, introrse to latrorse (rarely), expanded connective, usually with apiculus, [[Dehiscence (botany)|dehiscence]] along longitudinal tears. Hypogynous glands (0-)1-4, squamiform or elongated, fleshy, free or fused forming a lunate or annular [[nectar]]y over the receptacle. Superior gynoecium of 1(-2) apocarpous [[carpel]]s, sessile or stipitate (with a more or less elongated [[gynophore]]), sometimes not completely closed, [[Style (botany)|style]] usually developed, [[Stigma (botany)|stigma]] small or in the shape of a terminal or sub terminal disk or even lateral and oblique, often indented, papilous, moist or dry, [[ovule]]s 1-100 or more per carpel, anatropous, hemianatropous, amphitropous or orthotropous, mostly hemitropous, bitegmic, crassinucellate, [[chalaza]] with a ring of vascular bundles, the funiculus is occasionally absent and the ovule is fused to the placenta, marginal [[placentation]] with various dispositions or apical.
 
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* [[Fruit]] dehiscent or indehiscent, in [[achene]] or nucule, [[Follicle (fruit)|follicle]], [[drupe]] (with lignified endocarp) or falsely drupal (with lignified internal mesocarp), sometimes similar to a [[caryopsis]] as it is fused to the wall of the ovary and the testa, often lignified and serotinous; the fruit from the same inflorescence are sometimes fused forming a syncarp.
 
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* [[Seed]]s 1-many, sometimes winged, flat to rounded, with [[endosperm]] absent, present in ''Bellendina'', endotesta with an unusual layer containing crystals of [[calcium oxalate]] that is rarely absent, well differentiated [[embryo]], straight, dicotyledonous, but often with 3 or more (up to 9) large cotyledons, often auriculate.
 
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* [[Pollen]] in monads, triangular in polar view, (2-)3(-8)-aperturate, usually isopolar and triporate, biporate in ''[[Embothrium]]'' and the Banksieae tribe, colpoidate in ''[[Beauprea]]'', spherical in ''[[Aulax]]'' and ''[[Franklandia]]'' or strongly anisopolar in some species of ''[[Persoonia]]''; the openings of the former’s tetrads follow Garside’s Law.{{clarify|date=May 2014}}
 
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* [[Chromosome|Chromosomal number]]: ''n'' = 5, 7, 10-14, 26, 28; sizes range from very small (average of 1,0 μm) to very big (average of 14,4 μm) according to species; ''x'' = 7, 12.
 
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==Flowers==
 
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[[File:Pincushion hakea03.jpg|thumb|right|
 
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[[Inflorescence]] and leaves of the [[pin-cushion hakea]] (''Hakea laurina'')]]
 
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Generally speaking, the diagnostic feature of Proteaceae is the compound [[flower head]] or, more accurately, [[inflorescence]]. In many genera, the most obvious feature is the large and often very showy inflorescences, consisting of many small [[flower]]s densely packed into a compact head or spike. Even this character, however, does not occur in all Proteaceae; ''[[Adenanthos]]'' species, for example, have solitary flowers. In most Proteaceae species, the pollination mechanism is highly specialised. It usually involves the use of a "pollen-presenter", an area on the [[carpel|style]]-end that presents the [[pollen]] to the pollinator.<ref name="Watson 1992">{{cite web |author1=Watson, L.  |author2=Dallwitz, M. J.  |lastauthoramp=yes | title = Proteaceae | url = http://delta-intkey.com/angio/www/proteace.htm | year = 1992 onwards | work = [http://delta-intkey.com/angio/ The Families of Flowering Plants: Descriptions, Illustrations, Identification, Information retrieval] | date = 3 May 2006 | accessdate = 2006-06-26}}</ref>
 
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Proteaceae flower parts occur in fours, but the four [[tepal]]s are fused into a long, narrow tube with a closed cup at the top, and the filaments of the four [[stamen]]s are fused to the tepals, in such a way that the anthers are enclosed within the cup. The pistil initially passes along the inside of the perianth tube, so the stigma, too, is enclosed within the cup. As the flower develops, the pistil grows rapidly. Since the stigma is trapped, the style must bend to elongate, and eventually it bends so far, it splits the perianth along one seam. The style continues to grow until [[anthesis]], when the [[nectaries]] begin to produce [[nectar]]. At this time, the perianth splits into its component tepals, the cup splits apart, and the pistil is released to spring more or less upright.
 
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== Ecology ==
 
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[[File:Protea caffra IMG 2758.JPG|thumb|left|Inflorescence of ''Protea caffra'']]
 
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Many of the Proteaceae have specialised [[proteoid root]]s, masses of lateral roots and hairs forming a radial absorptive surface, produced in the leaf litter layer during seasonal growth, and usually shrivelling at the end of the growth season. They are an adaptation to growth in poor, phosphorus-deficient soils, greatly increasing the plants' access to scarce water and nutrients by exuding carboxylates that mobilise previously unavailable phosphorus. They also increase the root's absorption surface, but this is a minor feature, as it also increases competition for nutrients against its own root clusters.<ref name="Flora of Australia"/> However, this adaptation leaves them highly vulnerable to dieback caused by the ''[[Phytophthora cinnamomi]]'' [[water mould]], and generally intolerant of [[fertilizer|fertilization]]. Due to these specialized proteoid roots, the Proteaceae are one of few flowering plant families that do not form symbioses with [[arbuscular mycorrhiza]]l fungi. They exude large amounts of organic acids ([[citric acid]] and [[malic acid]]) every 2–3 days in order to aid the mobilization and absorption of phosphate. Many species are fire-adapted ([[pyrophyte]]s), meaning they have strategies for surviving fires that sweep through their habitat. Some are [[resprouter]]s, and have a thick rootstock buried in the ground that shoots up new stems after a fire, and others are [[reseeder]]s, meaning the adult plants are killed by the fire, but disperse their seeds, which are stimulated by the smoke to take root and grow. The heat was previously thought to have stimulated growth, but the chemicals in the smoke have now been shown to cause it.
 
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There are four dioecious genera (''[[Aulax]]'', ''[[Dilobeia]]'', ''[[Heliciopsis]]'' and ''[[Leucadendron]]''), 11 andromonoecious genera and some other genera have species that are cryptically andromonoecious: two species are sterile and only reproduce vegetatively (''Lomatia tasmanica'', ''Hakea pulvinifera''). The species vary between being autocompatible and autoincompatible, with intermediate situations; these situations sometimes occur in the same species. The flowers are usually protandrous. Just before anthesis, the anthers release their [[pollen]], depositing it onto the stigma, which in many cases has an enlarged fleshy area specifically for the deposition of its own pollen. Nectar-feeders are unlikely to come into contact with the anthers themselves, but can hardly avoid contacting the stigma; thus, the stigma functions as a [[pollen-presenter]], ensuring the nectar-feeders act as pollinators. The downside of this pollination strategy is that the probability of self-fertilisation is greatly increased; many Proteaceae counter this with strategies such as [[protandry]], self-incompatibility, or preferential abortion of selfed seed. The systems for presenting pollen are usually highly diverse, corresponding to the diversification of the pollinators. [[Pollination]] is carried out by [[bee]]s, [[beetle]]s, [[Fly|flies]], [[moth]]s, birds ([[honeyeater]]s, [[sunbird]]s, [[sugarbird]]s and [[hummingbird]]s) and mammals (rodents, small [[marsupial]]s, [[elephant shrew]]s and [[bats]]. The latter two means were evolutionarily derived from [[entomophily]] in different, independent events. The dispersion of some species exhibit the curious phenomenon of [[serotiny]], which is associated with their pyrophytic behaviour: these trees accumulate fruits on their branches whose outer layers or protective structures ([[bract]]s) are highly lignified and resistant to fire. The fruit only release their seeds when they have been burnt and when the ground has been fertilized with ashes from the fire and is free from competitors. Many species have seeds with [[elaiosome]]s that are dispersed by [[ant]]s; the seeds with wings or thistledown exhibit [[Seed dispersal syndrome#anemochory|anemochory]], while the drupes and other fleshy fruit exhibit [[Seed dispersal syndrome#zoochory|endozoochory]] as mammals and birds ingest them. Some African and Australian rodents are known to accumulate fruit and seeds of these plants in their nests in order to feed on them, although some manage to germinate.
 
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==Distribution ==
 
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Proteaceae are mainly a Southern Hemisphere family, with its main centres of diversity in Australia and South Africa. It also occurs in Central Africa, [[South America|South]] and [[Central America]], [[India]], eastern and south eastern [[Asia]], and [[Oceania]].<ref name="Flora of Australia">{{cite book | author = Orchard, Anthony E. (ed.) | title = Flora of Australia, Volume 16: Elaeagnaceae, Proteaceae 1 | url = http://www.deh.gov.au/biodiversity/abrs/online-resources/flora/main/ | chapter = Proteaceae | chapterurl = http://www.anbg.gov.au/abrs/online-resources/flora/stddisplay.xsql?pnid=1893 | location = Melbourne | publisher = Australian Biological Resources Study / CSIRO Publishing}}</ref> Only two species are known from New Zealand, although fossil pollen evidence suggests there were more previously.<ref>{{cite journal | author = Pole M| year = 1998 | title = The Proteaceae record in New Zealand| journal = Australian Systematic Botany | volume = 11 | issue = 4 | pages = 343–372 | doi = 10.1071/SB97019 }}</ref>
 
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It is a good example of a [[Gondwana]]n family, with taxa occurring on virtually every land mass considered a remnant of the ancient [[supercontinent]] Gondwana, except [[Antarctica]]. The family and subfamilies are thought to have diversified well before the fragmentation of Gondwana, implying all of them are well over 90 million years old. Evidence for this includes an abundance of proteaceous [[pollen]] found in the [[Cretaceous]] [[coal]] deposits of the [[South Island]] of [[New Zealand]]. It is thought to have achieved its present distribution largely by [[continental drift]] rather than dispersal across ocean gaps.<ref name="Weston 1996">{{cite book |author1=Weston, P. H.  |author2=Crisp, M. D.  |lastauthoramp=yes | year = 1996 | chapter = Trans-Pacific biogeographic patterns in the Proteaceae | editor = Keast, A. |editor2=Miller, S. E. | title = The origin and evolution of Pacific Island Biotas, New Guinea to eastern Polynesia: Patterns and processes | pages = 215–232 | location = Amsterdam | publisher=SPB Academic Publishing | isbn = 90-5103-136-X}}</ref>
 
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== Phytochemistry ==
 
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[[File:Brabejum stellatifolium seed.jpg|thumb|left|Fruit of ''Brabejum stellatifolium'']]
 
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No conclusive studies have been carried out on the chemical substances present in this broad family. The genera ''[[Protea]]'' and ''[[Faurea]]'' are unusual as they use [[xylose]] as the main sugar in their nectar and as they have high concentrations of polygalactol, while [[sucrose]] is the main sugar present in ''[[Grevillea]]''. [[Glycoside#Cyanogenic glycosides|Cyanogenic glycosides]], derived from [[tyrosine]], are often present, as are [[Condensed tannin|proanthocyanidine]]s ([[delphinidin]] and [[cyanidin]]), [[flavonol]]s ([[kaempferol]], [[quercetin]] and [[myricetin]]) and [[arbutin]]. [[Alkaloid]]s are usually absent. [[Iridoid]]s and [[ellagic acid]] are also absent. [[Saponin]]s and [[sapogenin]]s can be either present or absent in different species. Many species accumulate [[aluminium]].
 
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[[File:Leucadendron argenteum close.jpg|thumb|left|''[[Leucadendron argenteum]]'']]
 
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== Uses and cultivation ==
 
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[[File:MacNut2.JPG|thumb|right|Edible nuts of ''[[Macadamia]]'']]
 
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Many indigenous cultures have used Proteaceae as sustenance, medicine, for curing animal hides, as a source of dyes, firewood and as wood for construction. Aboriginal Australians eat the fruit of ''[[Persoonia]]'' and the seeds of species from other genera, including ''[[Gevuina]]'' and ''[[Macadamia]]'', form part of the diet not only of the indigenous peoples but they are also sold throughout the world. The tender shoots of ''[[Helicia]]'' species are used in Java and the nectar from the inflorescences of a number of species is drunk in Australia. Traditional medicines can be obtained from infusions of the roots, bark, leaves or flowers of many species that are used as topical applications for skin conditions or internally as tonics, aphrodisiacs, galactogens, to treat headaches, cough, dysentery, diarrhea, indigestion, stomach ulcers and kidney disease. The wood from the trees of this family are widely used in construction and for internal uses such as decoration, the wood from species of ''[[Protea]]'', ''[[Leucadendron]]'' and ''[[Grevillea]]'' are especially popular. Many species are used in gardening, particularly genera of ''[[Banksia]]'', ''[[Embothrium]]'', ''[[Grevillea]]'' and ''[[Waratah|Telopea]]''. Unfortunately this use has resulted in the introduction of exotic species that have become invasive, examples of this include the Hakea Willow (''[[Hakea salicifolia]]'') and the Silky Hakea (''[[Hakea sericea]]'') in Portugal. Two species of ''[[Macadamia]]'' are cultivated commercially for their edible nuts.
 
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''[[Gevuina avellana]]'' (Chilean hazel) is cultivated for its edible nuts in [[Chile]] and [[New Zealand]], and they are also used in the pharmaceutical industry for their humectant properties and as an ingredient in [[sunscreen]]s. It is the most resistant to cold of the tree families that produce nuts. It is also planted in the [[British Isles]] and on the Pacific coast of the [[United States]] for its tropical appearance and its ability to grow in [[Oceanic climate|cooler climates]] (it is also related to a common family in these latitudes).
 
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Many Proteaceae species are cultivated by the [[nursery (horticulture)|nursery]] industry, as barrier plants and for their prominent and distinctive flowers and foliage. Some species are of importance to the [[floristry|cut flower industry]], especially some ''[[Banksia]]'' and ''[[Protea]]'' species. Two species of the genus ''[[Macadamia]]'' are grown commercially for edible nuts. ''[[Gevuina avellana]]'' (Chilean hazelnut) tree is cultivated for its nuts in [[Chile]] and [[New Zealand]], which are edible, and are used in pharmaceutical industry for skin treatment because of their moisturizing properties and as ingredient in [[sunscreen]]s.
 
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Sugarbushes (''[[Protea]]''), pincushions (''[[Leucospermum]]'') and conebushes (''[[Leucadendron]]''), as well as others like pagodas (''[[Mimetes]]''), ''[[Aulax]]'' and blushing brides (''[[Serruria]]''), comprise one of the three main plant groups of [[fynbos]], which forms part of the [[Cape Floral Kingdom]], the smallest but richest plant kingdom for its size and the only kingdom contained within a single country. The other main groups of plants in fynbos are the [[Ericaceae]] and the [[Restionaceae]]. South African proteas are thus widely cultivated due to their many varied forms and unusual flowers. They are popular in South Africa for their beauty and their usefulness in [[wildlife garden]]s for attracting birds and useful insects.
 
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The species most valued as ornamentals are the trees that grow in southern latitudes as they give landscapes in [[temperate climates]] a tropical appearance; ''[[Lomatia ferruginea]]'' (Fuinque), ''[[Lomatia hirsuta]]'' (Radal) have been introduced in Western [[Europe]] and to the western [[United States]]. ''[[Embothrium coccineum]]'' (Chilean Firetree or ''Notro'') is highly valued in the British Isles for its dark red flowers and it can be found as far north as the [[Faroe Islands]] at a latitude of 62° north.
 
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Among the [[banksia]]s, many of which grow in temperate and Mediterranean climates, the vast majority are shrubs, only a few are trees that are valued for their height. Among the tallest species are: [[Banksia integrifolia|B.&nbsp;integrifolia]] with its subspecies [[Banksia integrifolia subsp. monticola|''B.&nbsp;integrifolia'' subsp. ''monticola'']], which is noteworthy as the plants that form the subspecies are the tallest trees of the banksias and they are the more frost-resistant than other banksias, [[Banksia seminuda|B.&nbsp;seminuda]], [[Banksia littoralis|B.&nbsp;littoralis]], [[Banksia serrata|B.&nbsp;serrata]]; among those that can be considered small trees or large shrubs: [[Banksia grandis|B.&nbsp;grandis]], [[Banksia prionotes|B.&nbsp;prionotes]], [[Banksia marginata|B.&nbsp;marginata]], [[Banksia coccinea|B.&nbsp;coccinea]] and [[Banksia speciosa|B.&nbsp;speciosa]]; all of these are planted in parks and gardens and even along roadsides because of their size. The rest of the species of this genus, around 170 species, are shrubs, although some of them are valued for their flowers.
 
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Another species that is cultivated in some parts of the world, although it is smaller, is ''[[Telopea speciosissima]]'' (Waratah), from the mountains of [[New South Wales]], [[Australia]].
 
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Some temperate climate species are cultivated more locally in Australia for their attractive appearance: ''[[Persoonia pinifolia]]'' (Pine –leaved [[Geebung]]) is highly valued for its vivid yellow flowers and grape-like fruit. ''[[Adenanthos sericeus]]'' (Woolly Bush) is planted for its attractive soft leaves and its small red or orange flowers. ''[[Hicksbeachia pinnatifolia]]'' ([[Beef]] [[Nut (fruit)|Nut]] or Red Bauple Nut) is commonly planted for its foliage and edible nuts.
 
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== Parasites ==
 
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[[File:Hakea purpurea.JPG|thumb|''Hakea purpurea'']]
 
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The Proteaceae are particularly susceptible to certain parasites, in particular the [[oomycete]] ''[[Phytophthora cinnamomi]]'', which causes severe root rot in the plants that grow in Mediterranean climates. ''[[Fusarium oxysporum]]'' causes a disease called fusariosis in roots that causes a yellowing and wilting, with serious ecological damages to woodland plants and economic losses in plants of commercial interest. Other common infections are caused by species of ''[[Botryosphaeria]]'', ''[[Rhizoctonia]]'', ''[[Armillaria]]'', ''[[Botryotinia|Botrytis]]'', ''[[Calonectria]]'' and other fungi.
 
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== Conservation status ==
 
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The [[IUCN]]<ref>IUCN 2006. 2006 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Downloaded on 22 February 2007</ref> considers that 47 Proteaceae species are threatened, of which one species, ''[[Stenocarpus dumbeensis]]'' [[André Guillaumin|Guillaumin]], 1935, from New Caledonia, is thought to be extinct. The species of this family are particularly susceptible to the destruction or fragmentation of their [[habitat]], fire, parasitic diseases, [[Competition (biology)|competition]] from introduced plants, [[soil]] degradation and other damage provoked by humans and their domesticated animals. The species are also affected by [[climate change]].
 
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== Fossils ==
 
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[[File:Lambertia multiflora.jpg|thumb|''Lambertia multiflora'']]
 
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The proteaceae have a rich fossil record, despite the inherent difficulties in identifying remains that do not show diagnostic characteristics. Identification usually comes from using a combination of brachy-paracytic stomata and the unusual [[trichome]] bases or, in other cases, the unusual structure of pollen tetrads. Fossils attributable to this family have been found on the majority of areas that formed the [[Gondwana]] supercontinent. A wide variety of pollen belonging to this family dating back to the Upper [[Cretaceous]] ([[Campanian]]-[[Maastrichtian]]) from the south east of Australia and pollen from the Middle Cretaceous ([[Cenomanian]]-[[Turonian]]) from northern Africa and Peru described as ''Triorites africaensis''. The first macrofossils appear twenty million years later in the [[Palaeocene]] of South America and the north east of Australia. The fossil record of some areas, such as New Zealand and Tasmania, show a greater biodiversity for Proteaceae than currently exists, which supports the fact that the distribution of many taxa has changed drastically with the passage of time and that the family has suffered a general decline, including high levels of extinction during the [[Cenozoic]].
 
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==Taxonomy==
 
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First described by French botanist [[Antoine Laurent de Jussieu]], the family Proteaceae is a fairly large one, with around 80 genera, but less than 2000 species. It is recognised by virtually all [[Taxonomy (biology)|taxonomists]]. Firmly established under classical [[Linnaean taxonomy]], it is also recognised by the [[cladistics]]-based [[APG system|APG]] and [[APG II system|APG II]] systems. It is placed in the order [[Proteales]], whose placement has itself varied.
 
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The framework for classification of the genera within Proteaceae was laid by [[Lawrence Alexander Sidney Johnson|Lawrie Johnson]] and [[Barbara G. Briggs|Barbara Briggs]] in their influential 1975 monograph "[[On the Proteaceae: the evolution and classification of a southern family]]".<ref name="Lawrence 1975">{{cite journal|author = [[Lawrence Alexander Sidney Johnson|L. A. S. Johnson]] and [[Barbara G. Briggs|Briggs, B. G.]] | year = 1975 | title = On the Proteaceae: the evolution and classification of a southern family | journal = Journal of the Linnean Society of London. Botany | volume = 70 | pages = 83–182 | doi=10.1111/j.1095-8339.1975.tb01644.x|issue = 2}}</ref> Their classification has been refined somewhat over the ensuing three decades, most notably by Peter H. Weston and Nigel Barker in 2006. Proteaceae are now divided into five subfamilies: [[Bellendenoideae]], [[Persoonioideae]], [[Symphionematoideae]], [[Proteoideae]] and [[Grevilleoideae]].<ref name="Weston 2006">{{cite journal |author1=Weston, Peter H. |author2=Barker, Nigel P. | year = 2006 | title = A new suprageneric classification of the Proteaceae, with an annotated checklist of genera | journal = Telopea | volume = 11 | issue = 3 | pages = 314–344|doi=10.7751/telopea20065733}}</ref> In 2008 Mast and colleagues updated ''Macadamia'' and related genera in tribe Macadamieae. Furthermore, ''Orites megacarpus'' was found as polyphyletic with and not in the ''Orites'' genus, nor in the tribe Roupaleae, instead in the tribe Macadamieae also, hence given the new genus ''Nothorites'' and species name combination of ''Nothorites megacarpus''.<ref name=Mastetal2008>{{Cite journal
 
−
| last1 = Mast | first1 = Austin R.
 
−
| last2 = Willis | first2 = Crystal L.
 
−
| last3 = Jones | first3 = Eric H.
 
−
| last4 = Downs | first4 = Katherine M.
 
−
| last5 = Weston | first5 = Peter H.
 
−
|date=July 2008
 
−
| title = A smaller ''Macadamia'' from a more vagile tribe: inference of phylogenetic relationships, divergence times, and diaspore evolution in ''Macadamia'' and relatives (tribe Macadamieae; Proteaceae)
 
−
| journal = American Journal of Botany
 
−
| volume = 95 | issue = 7 | pages = 843–870
 
−
| issn = 1537-2197
 
−
| doi = 10.3732/ajb.0700006
 
−
| url = http://www.amjbot.org/content/95/7/843
 
−
| accessdate = 4 Apr 2013 | pmid=21632410
 
−
}}</ref> The full arrangement, according to Weston and Barker (2006) with the updates to genera from Mast et al. (2008), is as follows:
 
−
[[File:Banksia coccinea (Illustrationes Florae Novae Hollandiae plate 3).jpg|thumb|right|Flowers, leaves and fruit of ''Banksia coccinea'', from [[Ferdinand Bauer]]'s 1813 flora ''[[Illustrationes Florae Novae Hollandiae]]'']]
 
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:Family Proteaceae
 
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::Subfamily [[Bellendenoideae]]
 
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:::::''[[Bellendena]]''
 
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::Subfamily [[Persoonioideae]]
 
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:::Tribe [[Placospermeae]]
 
−
:::::''[[Placospermum]]''
 
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:::Tribe [[Persoonieae]]
 
−
:::::''[[Toronia]]''&nbsp;— ''[[Garnieria]]''&nbsp;— ''[[Acidonia]]''&nbsp;— ''[[Persoonia]]''
 
−
::Subfamily [[Symphionematoideae]]
 
−
:::::''[[Agastachys odorata|Agastachys]]''&nbsp;— ''[[Symphionema]]''
 
−
::Subfamily [[Proteoideae]]
 
−
:::[[incertae sedis]]
 
−
:::::''[[Eidothea]]''&nbsp;— ''[[Beauprea]]''&nbsp;— ''[[Beaupreopsis]]''&nbsp;— ''[[Dilobeia]]''&nbsp;— ''[[Cenarrhenes]]''&nbsp;— ''[[Franklandia]]''
 
−
:::Tribe [[Conospermeae]]
 
−
::::Subtribe [[Stirlingiinae]]
 
−
:::::''[[Stirlingia]]''
 
−
::::Subtribe [[Conosperminae]]
 
−
:::::''[[Conospermum]]''&nbsp;— ''[[Synaphea]]''
 
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:::Tribe [[Petrophileae]]
 
−
:::::''[[Petrophile]]''&nbsp;— ''[[Aulax]]''
 
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:::Tribe [[Proteeae]]
 
−
:::::''[[Protea]]''&nbsp;— ''[[Faurea]]''
 
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:::Tribe [[Leucadendreae]]
 
−
::::Subtribe [[Isopogoninae]]
 
−
:::::''[[Isopogon]]''
 
−
::::Subtribe [[Adenanthinae]]
 
−
:::::''[[Adenanthos]]''
 
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::::Subtribe [[Leucadendrinae]]
 
−
:::::''[[Leucadendron]]''&nbsp;— ''[[Serruria]]''&nbsp;— ''[[Paranomus]]''&nbsp;— ''[[Vexatorella]]''&nbsp;— ''[[Sorocephalus]]''&nbsp;— ''[[Spatalla]]''&nbsp;— ''[[Leucospermum]]''&nbsp;— ''[[Mimetes]]''&nbsp;— ''[[Diastella]]''&nbsp;— ''[[Orothamnus]]''
 
−
::Subfamily [[Grevilleoideae]]
 
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:::[[incertae sedis]]
 
−
:::::''[[Sphalmium]]''&nbsp;— ''[[Carnarvonia]]''
 
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:::Tribe [[Roupaleae]]
 
−
::::[[incertae sedis]]
 
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:::::''[[Megahertzia]]''&nbsp;— ''[[Knightia (plant)|Knightia]]''&nbsp;— ''[[Eucarpha]]''&nbsp;— ''[[Triunia]]''
 
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::::Subtribe [[Roupalinae]]
 
−
:::::''[[Roupala]]''&nbsp;— ''[[Neorites]]''&nbsp;— ''[[Orites]]''
 
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::::Subtribe [[Lambertiinae]]
 
−
:::::''[[Lambertia]]''&nbsp;— ''[[Xylomelum]]''
 
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::::Subtribe [[Heliciinae]]
 
−
:::::''[[Helicia]]''&nbsp;— ''[[Hollandaea]]''
 
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::::Subtribe [[Floydiinae]]
 
−
:::::''[[Darlingia]]''&nbsp;— ''[[Floydia]]''
 
−
:::Tribe [[Banksieae]]
 
−
::::Subtribe [[Musgraveinae]]
 
−
:::::''[[Musgravea]]''&nbsp;— ''[[Austromuellera]]''
 
−
::::Subtribe [[Banksiinae]]
 
−
:::::''[[Banksia]]''&nbsp;— ''[[Dryandra]]''
 
−
:::Tribe [[Embothrieae]]
 
−
::::Subtribe [[Lomatiinae]]
 
−
:::::''[[Lomatia]]''
 
−
::::Subtribe [[Embothriinae]]
 
−
:::::''[[Embothrium]]''&nbsp;— ''[[Oreocallis]]''&nbsp;— ''[[Alloxylon]]''&nbsp;— ''[[Telopea (genus)|Telopea]]''
 
−
::::Subtribe [[Stenocarpinae]]
 
−
:::::''[[Stenocarpus]]''&nbsp;— ''[[Strangea]]''
 
−
::::Subtribe [[Hakeinae]]
 
−
:::::''[[Opisthiolepis]]''&nbsp;— ''[[Buckinghamia]]''&nbsp;— ''[[Hakea]]''&nbsp;— ''[[Grevillea]]''&nbsp;— ''[[Finschia]]''
 
−
:::Tribe [[Macadamieae]]
 
−
::::Subtribe [[Macadamiinae]]
 
−
:::::''[[Macadamia]]''&nbsp;— ''[[Lasjia]]''&nbsp;— ''[[Nothorites]]''&nbsp;— ''[[Panopsis]]''&nbsp;— ''[[Brabejum]]''
 
−
::::Subtribe [[Malagasiinae]]
 
−
:::::''[[Malagasia]]''&nbsp;— ''[[Catalepidia]]''
 
−
::::Subtribe [[Virotiinae]]
 
−
:::::''[[Virotia]]''&nbsp;— ''[[Athertonia]]''&nbsp;— ''[[Heliciopsis]]''
 
−
::::Subtribe [[Gevuininae]]
 
−
:::::''[[Cardwellia]]''&nbsp;— ''[[Sleumerodendron]]''&nbsp;— ''[[Euplassa]]''&nbsp;— ''[[Gevuina]]''&nbsp;— ''[[Bleasdalea]]''&nbsp;— ''[[Hicksbeachia]]''&nbsp;— ''[[Kermadecia]]''&nbsp;— ''[[Turrillia]]''
 
−
 
 
−
== References ==
 
−
[[File:Woorikee2000.jpg|thumb|''Isopogon anemonifolius'']]
 
−
{{Reflist | 27em}}
 
−
 
 
−
* {{cite book
 
−
| author = Weston, P.H.
 
−
| chapter = Proteaceae
 
−
| title = Kubitzki, K. (Editor). The Families and Genera of Vascular Plants. IX. Flowering Plants - Eudicots.
 
−
| year = 2007
 
−
| publisher = Springer-Verlag: Berlín
 
−
| isbn = 3-540-32214-0
 
−
}}
 
−
* {{cite journal
 
−
|author1=Hoot, S.B.  |author2=Douglas, A.W.
 
−
|lastauthoramp=yes | title = Phylogeny of the Proteaceae based on ''atp''B and ''atp''B-''rbc''L intergenic spacer region sequences
 
−
| year = 1998
 
−
| journal = Australian Systematic Botany
 
−
| volume = 11
 
−
| number =
 
−
| pages=301–320
 
−
| doi = 10.1071/sb98027
 
−
}}
 
−
* {{Cite web|author1=Watson, L. |author2=Dallwitz, M.J.
 
−
|lastauthoramp=yes |title= The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 29th July 2006.
 
−
|year= 1992
 
−
|url= http://delta-intkey.com
 
−
|accessdate= 31 January 2007
 
−
}}
 
−
 
 
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== External links ==
 
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{{wikispecies}}
 
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{{commons category|Proteaceae}}
 
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* [http://www.anbg.gov.au/images/photo_cd/proteaceae/ Images of Proteaceae from the Australian National Botanical Gardens]
 
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* [http://www.mobot.org/MOBOT/Research/APweb/maps/proteaceaemap.gif Map]
 
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{{taxonbar}}
 
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{{Authority control}}
 
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[[Category:Proteaceae| ]]
 
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[[Category:Eudicot families]]
 
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[[Category:Extant Late Cretaceous first appearances]]
 

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