The Proteaceae /ˌproʊtiˈeɪsiː/ form a family of owering plants predominantly distributed in the

Southern Hemisphere. The family comprises 83 genera with about 1,660 known species.[3] Australia
and South Africa have the greatest concentrations of diversity. Together with the Platanaceae (plane
trees), Nelumbonaceae (the sacred lotus) and in the recent APG IV system the Sabiaceae, they make
up the order Proteales. Well-known Proteaceae genera include Protea, Banksia, Embothrium,
Grevillea, Hakea, 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 owers. The nuts of Macadamia integrifolia are widely grown
commercially and consumed, as are those of Gevuina avellana on a smaller scale.

Etymology
The name Proteaceae was adapted by Robert Brown from the name Proteae coined in 1789 for the
family by Antoine Laurent de Jussieu, based on the genus Protea, which in 1767, Carl Linnaeus
derived from the name of the Greek god Proteus, a deity who was able to change between many
forms.[4][5] This is an appropriate image, seeing as the family is known for its astonishing variety and
diversity of owers and leaves.[citation needed]

Description

Rhopala heterophylla
The genera of Proteaceae are highly varied, with Banksia in particular providing a striking example
of adaptive radiation in plants.[6] This variability makes it impossible to provide a simple, diagnostic
identi cation key for the family, although individual genera may be easily identi ed.

• Proteaceae range from prostrate shrubs to tall forest trees, of 40 m in height, and are usually
of medium height or low or perennial shrubs, except for some Stirlingia species that are
herbs. Some species 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 bi d.
• Leaves rarely aromatic, usually alternate, and in a spiral, rarely opposed, or verticilate;
coriaceous, rarely eshy 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 frequently with a swollen base but rarely sheathed (sometimes
in Synaphea), without stipules; pinnate sometimes palmate or parallel 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 idioblasts, rare
secretory cavities. Brachy-paracytic stomata (laterocytic in Bellendena).
Plant stems with two types of radii, wide and multi-serrated or narrow and uni-serrated, phloem
strati ed or not, trilacunar nodes with three leaf traces (rarely unilacunar with one trace), sclereids
frequent; bark with lenticels frequently horizontally enlarged, cork cambium present, usually
super cial. Roots lateral and short, often grouped in bundles (proteoid roots) with very dense root
hairs, rarely with mycorrhiza.

• Plants usually hermaphroditic, more rarely monoecious, dioecious or andromonoecious.

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 • In orescences very variable, simple or compound, axillary or terminal, lateral owers
solitary or in pairs, rarely with a terminal ower, racemiform, paniculate or condensed,
usually with bracts, sometimes converted into leaves or squamiform, forming a type of cone,
or with bright colours, forming an involucre or pseudanthium, the peduncles and pedicels
sometimes contracted, compacted with the rachis, in some cases the congested
in orescences form super in orescences (some Alloxylon); very rarely the owers are
solitary and axillary near the end of branches; in species with lignotubers the owers
sometimes grow from these and pass through the soil (geophytes).
• Flowers 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) tepals (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) stamens, all fertile or some
converted into staminodes, usually lamentous, laments partially or totally fused to the
tepals, rarely free, basi xed anthers adnate, ditheous, tetrasporangiate, sometimes unilocular
and bisporagiate, introrse to latrorse (rarely), expanded connective, usually with apiculus,
dehiscence along longitudinal tears. Hypogynous glands (0-)1–4, squamiform or elongated,
eshy, free or fused forming a lunate or annular nectary over the receptacle. Superior
gynoecium of 1(−2) apocarpous carpels, sessile or stipitate (with a more or less elongated
gynophore), sometimes not completely closed, style usually developed, stigma small or in
the shape of a terminal or sub terminal disk or even lateral and oblique, often indented,
papilous, moist or dry, ovules 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.
• Fruit dehiscent or indehiscent, in achene or nucule, follicle, drupe (with ligni ed endocarp)
or falsely drupal (with ligni ed internal mesocarp), sometimes similar to a caryopsis as it is
fused to the wall of the ovary and the testa, often ligni ed and serotinous; the fruit from the
same in orescence are sometimes fused forming a syncarp.
• Seeds 1-many, sometimes winged, at 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.
• Pollen in monads, triangular in polar view, (2-)3(−8)-aperturate, usually isopolar and
triporate, biporate in Embothrium and the tribe Banksieae, 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.[clari cation needed]
• 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.
Flowers

In orescence and leaves of the pin-cushion hakea (Hakea laurina)

Generally speaking, the diagnostic feature of Proteaceae is the compound ower head or, more
accurately, in orescence. In many genera, the most obvious feature is the large and often very
showy in orescences, consisting of many small owers densely packed into a compact head or
spike. Even this character, however, does not occur in all Proteaceae; Adenanthos species, for
example, have solitary owers. In most Proteaceae species, the pollination mechanism is highly
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 specialised. It usually involves the use of a "pollen-presenter", an area on the style-end that presents
the pollen to the pollinator.[7]

Proteaceae ower parts occur in fours, but the four tepals are fused into a long, narrow tube with a
closed cup at the top, and the laments of the four stamens 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 ower 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.

Ecology

In orescence of Protea caffra

Many of the Proteaceae have specialised proteoid roots, 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-
de cient 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.[8] However, this adaptation leaves them highly vulnerable to dieback caused by
the Phytophthora cinnamomi water mould, and generally intolerant of fertilization. Due to these
specialized proteoid roots, the Proteaceae are one of few owering plant families that do not form
symbioses with arbuscular mycorrhizal 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 re-adapted (pyrophytes), meaning they have strategies for surviving res that
sweep through their habitat. Some are resprouters, and have a thick rootstock buried in the ground
that shoots up new stems after a re, and others are reseeders, meaning the adult plants are killed by
the re, 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.

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 owers are usually protandrous. Just
before anthesis, the anthers release their pollen, depositing it onto the stigma, which in many cases
has an enlarged eshy area speci cally 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 diversi cation of the pollinators. Pollination is carried
out by bees, beetles, ies, moths, birds (honeyeaters, sunbirds, sugarbirds and hummingbirds) and
mammals (rodents, small marsupials, elephant shrews 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
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behaviour: these trees accumulate fruits on their branches whose outer layers or protective
structures (bracts) are highly ligni ed and resistant to re. The fruit only release their seeds when
they have been burnt and when the ground has been fertilized with ashes from the re and is free
from competitors. Many species have seeds with elaiosomes that are dispersed by ants; the seeds
with wings or thistledown exhibit anemochory, while the drupes and other eshy fruit exhibit
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.

Distribution
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 and Central America, India, eastern and
south eastern Asia, and Oceania.[8] Only two species are known from New Zealand, although fossil
pollen evidence suggests there were more previously.[9]

It is a good example of a Gondwanan 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 diversi ed 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.[10]

Phytochemistry

Fruit of Brabejum stellatifolium

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. Cyanogenic glycosides, derived from tyrosine, are often present, as are
proanthocyanidines (delphinidin and cyanidin), avonols (kaempferol, quercetin and myricetin) and
arbutin. Alkaloids are usually absent. Iridoids and ellagic acid are also absent. Saponins and
sapogenins can be either present or absent in different species. Many species accumulate
aluminium.

Leucadendron argenteum
Uses and cultivation

Edible nuts of Macadamia

Many traditional cultures have used Proteaceae as sustenance, medicine, for curing animal hides, as
a source of dyes, rewood 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 of the indigenous peoples but are also sold throughout the world. The tender shoots
of Helicia species are used in Java, and the nectar from the in orescences of a number of species is
drunk in Australia. Traditional medicines can be obtained from infusions of the roots, bark, leaves,
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 or owers of many species that are used as topical applications for skin conditions or internally as
tonics, aphrodisiacs, and galactogens to treat headaches, cough, dysentery, diarrhea, indigestion,
stomach ulcers, and kidney disease. The wood from the trees of this family is widely used in
construction and for internal uses such as decoration; the wood from species of Protea,
Leucadendron and Grevillea is especially popular. Many species are used in gardening, particularly
genera of Banksia, Embothrium, Grevillea, and Telopea. This use has resulted in the introduction of
exotic species that have become invasive; examples 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. Gevuina avellana
(Chilean hazel) is also 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
sunscreens. It is the most cold-resistant of the tree families that produce nuts.[citation needed] It is also
planted in the British Isles and on the Paci c coast of the United States for its tropical appearance
and its ability to grow in cooler climates.

Many Proteaceae species are cultivated by the nursery industry as barrier plants and for their
prominent and distinctive owers and foliage. Some species are of importance to the cut ower
industry, especially some Banksia and Protea species. Two species of the genus Macadamia are
grown commercially for edible nuts.

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 owers. They are popular in South Africa for
their beauty and their usefulness in wildlife gardens for attracting birds and useful insects.

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 owers and can be found as far north as the Faroe Islands at a latitude of 62° north.

Among the banksias, 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: B. integrifolia with its subspecies B. 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, B. seminuda, B. littoralis, B. serrata; among those that can be
considered small trees or large shrubs: B. grandis, B. prionotes, B. marginata, B. coccinea and
B. 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 owers.

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.

Some temperate climate species are cultivated more locally in Australia for their attractive
appearance: Persoonia pinifolia (pine-leaved geebung) is valued for its vivid yellow owers and
grape-like fruit. Adenanthos sericeus (woolly bush) is planted for its attractive soft leaves and its
small red or orange owers. Hicksbeachia pinnatifolia (beef nut, red bauple nut) is commonly
planted for its foliage and edible nuts.
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Parasites

Hakea purpurea
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, Botrytis, Calonectria and other fungi.

Conservation status
The IUCN[11] considers that 47 Proteaceae species are threatened, of which one species, Stenocarpus
dumbeensis 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, re, parasitic
diseases, competition from introduced plants, soil degradation and other damage provoked by
humans and their domesticated animals. The species are also affected by climate change.

Fossils

Lambertia multi ora

The Proteaceae have a rich fossil record, despite the inherent dif culties in identifying remains that
do not show diagnostic characteristics. Identi cation 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.[citation needed] Xylocaryon was identi ed as a member of the Proteaceae from the
similarity of its fruit to the extant genus Eidothea.[12] 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 rst 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.[citation needed]

Taxonomy

Isopogon anemonifolius
First described by French botanist Antoine Laurent de Jussieu, the family Proteaceae is a fairly large
one, with around 80 genera, but less than 2,000 species. It is recognised by virtually all taxonomists.
Firmly established under classical Linnaean taxonomy, it is also recognised by the cladistics-based
APG and APG II systems. It is placed in the order Proteales, whose placement has itself varied.
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 A classi cation of the genera within Proteaceae was made by Lawrie Johnson and Barbara Briggs[13]
in their in uential 1975 monograph "On the Proteaceae: the evolution and classi cation of a
southern family",[14] until it was largely superseded by the molecular studies of Peter H. Weston and
Nigel Barker in 2006. Proteaceae are now divided into ve subfamilies: Bellendenoideae,
Persoonioideae, Symphionematoideae, Proteoideae and Grevilleoideae.[14] In 2008 Mast and
colleagues updated Macadamia and related genera in tribe Macadamieae. Furthermore, Orites
megacarpus was found not to be within the genus Orites, nor in the tribe Roupaleae, instead in the
tribe Macadamieae, hence given the new species name Nothorites megacarpus.[15] The full
arrangement, according to Weston and Barker (2006) with the updates to genera from Mast et al.
(2008), is as follows:

Flowers, leaves and fruit of Banksia coccinea, from Ferdinand Bauer's 1813 ora Illustrationes
Florae Novae Hollandiae
Family Proteaceae
Subfamily Bellendenoideae
Bellendena
Subfamily Persoonioideae
Tribe Placospermeae
Placospermum
Tribe Persoonieae
Persoonia
Subfamily Symphionematoideae
Agastachys — Symphionema
Subfamily Proteoideae
incertae sedis
Eidothea — Beauprea — Beaupreopsis — Dilobeia — Cenarrhenes — Franklandia
Tribe Conospermeae
Subtribe Stirlingiinae
Stirlingia
Subtribe Conosperminae
Conospermum — Synaphea
Tribe Petrophileae
Petrophile — Aulax
Tribe Proteeae
Protea — Faurea
Tribe Leucadendreae
Subtribe Isopogoninae
Isopogon
Subtribe Adenanthinae
Adenanthos
Subtribe Leucadendrinae
Leucadendron — Serruria — Paranomus — Vexatorella — Sorocephalus — Spatalla —
Leucospermum — Mimetes — Diastella — Orothamnus
Subfamily Grevilleoideae
incertae sedis
Sphalmium — Carnarvonia
Tribe Roupaleae
incertae sedis
Megahertzia — Knightia — Eucarpha — Triunia
Subtribe Roupalinae
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Roupala — Neorites — Orites
Subtribe Lambertiinae
Lambertia — Xylomelum
Subtribe Heliciinae
Helicia — Hollandaea
Subtribe Floydiinae
Darlingia — Floydia
Tribe Banksieae
Subtribe Musgraveinae
Musgravea — Austromuellera
Subtribe Banksiinae
Banksia
Tribe Embothrieae
Subtribe Lomatiinae
Lomatia
Subtribe Embothriinae
Embothrium — Oreocallis — Alloxylon — Telopea
Subtribe Stenocarpinae
Stenocarpus — Strangea
Subtribe Hakeinae
Opisthiolepis — Buckinghamia — Hakea — Grevillea — Finschia
Tribe Macadamieae
Subtribe Macadamiinae
Macadamia — Lasjia — Nothorites — Panopsis — Brabejum
Subtribe Malagasiinae
Malagasia — Catalepidia
Subtribe Virotiinae
Virotia — Athertonia — Heliciopsis
Subtribe Gevuininae
Cardwellia — Euplassa — Gevuina — Bleasdalea — Hicksbeachia — Kermadecia
References
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j.1095-8339.1975.tb01644.x.

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(July 2008). "A smaller Macadamia from a more vagile tribe: inference of phylogenetic
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Macadamieae; Proteaceae)". American Journal of Botany. 95 (7): 843–870. doi:10.3732/
ajb.0700006. ISSN 1537-2197. PMID 21632410.
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Vascular Plants. IX. Flowering Plants – Eudicots. Springer-Verlag: Berlín.
ISBN 978-3-540-32214-6.
• Hoot, S.B. & Douglas, A.W. (1998). "Phylogeny of the Proteaceae based on atpB and atpB-
rbcL intergenic spacer region sequences". Australian Systematic Botany. 11 (4): 301–320.
doi:10.1071/sb98027.
• Ramsey, H.P.. (1963). "Chromosome numbers in the proteaceae". Australian Journal of
Botany. 11 (1): 1–20. doi:10.1071/BT9630001.
• Watson, L. & Dallwitz, M.J. (1992). "The families of owering plants: descriptions,
illustrations, identi cation, and information retrieval. Version: 29th July 2006". Retrieved
31 January 2007.
• Brown, R. On the Proteaceae of Jussieu Proceedings of the Linnean Society 10:15-216.
External links
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 • Images of Proteaceae from the Australian National Botanical Gardens
• Map

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Genera of Proteaceae

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Families of owering plants (APG IV)

Taxon identi ers

Proteaceae • Wikidata: Q157228
• Wikispecies: Proteaceae
• APNI: 54427
• ATRF: Proteaceae
• BOLD: 148506
• CoL: F4J
• EoL: 4370
• EPPO: 1PROF
• FloraBase: 22793
• FoAO2: Proteaceae
• FoC: 10730
• GBIF: 2414
• GRIN: 922
• iNaturalist: 64517
• IPNI: 30004394-2
• IRMNG: 114453
• ITIS: 27781
• NCBI: 4328
• NZOR:
11f0fb59-1293-42fe-8c8f-87ad5b72d3cc
• Open Tree of Life: 209175
• Paleobiology Database: 55681
• POWO: urn:lsid:ipni.org:names:30004394-2
• Tropicos: 42000186
• VicFlora: f48162c2-e198-4065-
a64c-3d1081392043
• Watson & Dallwitz: proteace
• WFO: wfo-7000000499
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Authority • Israel
control • Latvia
databases: • Japan
National

Categories:
• Proteaceae
• Eudicot families
• Extant Cenomanian rst appearances
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