The Proteaceae form a
family of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s predominantly distributed in the
Southern Hemisphere. The family comprises 83
genera with about 1,660 known
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 appropriat ...
.
Together with the
Platanaceae and
Nelumbonaceae, they make up the order
Proteales. Well-known genera include ''
Protea'', ''
Banksia'', ''
Embothrium'', ''
Grevillea'', ''
Hakea'' and ''
Macadamia''. Species such as the New South Wales waratah (''
Telopea speciosissima
''Telopea speciosissima'', commonly known as the New South Wales waratah or simply waratah, is a large shrub in the plant family Proteaceae. It is endemic to New South Wales in Australia and is the floral emblem of that state. No subspecies ...
''), king protea (''
Protea cynaroides''), and various species of ''Banksia'', ''soman'', and ''Leucadendron'' are popular
cut flowers. The nuts of ''
Macadamia integrifolia'' are widely grown commercially and consumed, as are those of
Gevuina avellana on a smaller scale.
Australia
Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
and
South Africa
South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
have the greatest concentrations of diversity.
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
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
derived from the name of the Greek god
Proteus
In Greek mythology, Proteus (; Ancient Greek: Πρωτεύς, ''Prōteus'') is an early prophetic sea-god or god of rivers and oceanic bodies of water, one of several deities whom Homer calls the "Old Man of the Sea" ''(hálios gérôn)''. ...
, a deity who 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.
Description
The genera of Proteaceae are highly varied, with ''
Banksia'' in particular providing a striking example of
adaptive radiation
In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic in ...
in plants.
This variability makes it impossible to provide a simple, diagnostic identification key for the family, although individual genera may be easily identified.
* 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
''Embothrium coccineum'', Chilean firetree or Chilean firebush, commonly known in Chile and Argentina as ''notro'', ''ciruelillo'' and ''fósforo'' is a small evergreen tree in the flowering plant family Proteaceae. It grows in the temperate for ...
''), 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.
*
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 frequently with a swollen base but rarely sheathed (sometimes in ''
Synaphea
''Synaphea'' is a genus of small shrubs and is Endemism, endemic to Western Australia. Synapheas have variably shaped leaves but consistently yellow flowers with an unusual pollination mechanism.
Description
Plants in the genus ''Synaphea'' are ...
''), 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
''Bellendena montana'', commonly known as mountain rocket, is a species of low-growing multi-stemmed shrub in the plant family Proteaceae. It is endemic to high-altitude subalpine and alpine regions in Tasmania, Australia. The prominent white fl ...
'').
Plant stems with two types of radii, wide and multi-serrated or narrow and uni-serrated,
phloem stratified or not, trilacunar nodes with three
leaf traces (rarely unilacunar with one trace),
sclereids frequent;
bark
Bark may refer to:
* Bark (botany), an outer layer of a woody plant such as a tree or stick
* Bark (sound), a vocalization of some animals (which is commonly the dog)
Places
* Bark, Germany
* Bark, Warmian-Masurian Voivodeship, Poland
Arts, e ...
with
lenticels frequently horizontally enlarged,
cork cambium present, usually superficial.
Roots lateral and short, often grouped in bundles (''proteoid roots'') with very dense root hairs, rarely with
mycorrhiza.
*
Plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s usually hermaphroditic, more rarely monoecious, dioecious or andromonoecious.
*
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 forme ...
s very variable, simple or compound, axillary or terminal, lateral flowers solitary or in pairs, rarely with a terminal flower,
racemiform,
paniculate
A panicle is a much-branched inflorescence. (softcover ). Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers (and fruit) be pedicellate (having a single stem per flower). The branches of a panicle are ...
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 inflorescences form super inflorescences (some ''
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 ...
''); 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).
*
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
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)
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
staminode
In botany, a staminode is an often rudimentary, sterile or abortive stamen, which means that it does not produce pollen.Jackson, Benjamin, Daydon; ''A Glossary of Botanic Terms with their Derivation and Accent''; Published by Gerald Duckworth & C ...
s, usually filamentous, filaments partially or totally fused to the tepals, rarely free, basifixed
anthers adnate, ditheous, tetrasporangiate, sometimes
unilocular
A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus).
In angiosperms (flowering plants), the term ''locule'' usu ...
and bisporagiate, introrse to latrorse (rarely), expanded connective, usually with apiculus,
dehiscence along longitudinal tears. Hypogynous glands (0-)1-4, squamiform or elongated, fleshy, 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
Placentation refers to the formation, type and structure, or arrangement of the placenta. The function of placentation is to transfer nutrients, respiratory gases, and water from maternal tissue to a growing embryo, and in some instances to remo ...
with various dispositions or apical.
*
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 partic ...
dehiscent or indehiscent, in
achene or nucule,
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.
*
Seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
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.
*
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
'' Aulax'' is a South African Proteaceae genus of just three species of evergreen shrubs commonly known as "featherbushes". The name ''Aulax'' was named by a botanist named Linnaeus; he named ''Aulax'' after the Greek god Proteus, who is known to ...
'' and ''
Franklandia'' or strongly anisopolar in some species of ''
Persoonia''; the openings of the former's tetrads follow Garside's Law.
*
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
Generally speaking, the diagnostic feature of Proteaceae is the compound
flower head or, more accurately,
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 forme ...
. In many genera, the most obvious feature is the large and often very showy inflorescences, consisting of many small
flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s densely packed into a compact head or spike. Even this character, however, does not occur in all Proteaceae; ''
Adenanthos
''Adenanthos'' is a genus of Australian native shrubs in the flowering plant family Proteaceae. Variable in habit and leaf shape, it is the only genus in the family where solitary flowers are the norm. It was discovered in 1791, and formally publ ...
'' 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
style-end that presents the
pollen to the pollinator.
Proteaceae flower parts occur in fours, but the four
tepals are fused into a long, narrow tube with a closed cup at the top, and the filaments 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 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
Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period.
The onset of anthesis is spectacular in some species. In ''Banksia'' species, for example, anthesis involves the extension ...
, 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
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-deficient soils, greatly increasing the plants' access to scarce water and nutrients by exuding
carboxylates
In organic chemistry, a carboxylate is the conjugate base of a carboxylic acid, (or ). It is an ion with negative charge.
Carboxylate salts are salts that have the general formula , where M is a metal and ''n'' is 1, 2,...; ''carboxyla ...
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.
However, this adaptation leaves them highly vulnerable to dieback caused by the ''
Phytophthora cinnamomi''
water mould, and generally intolerant of
fertilization
Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
. Due to these specialized proteoid roots, the Proteaceae are one of few flowering plant families that do not form symbioses with
arbuscular mycorrhizal fungi. They exude large amounts of organic acids (
citric acid
Citric acid is an organic compound with the chemical formula HOC(CO2H)(CH2CO2H)2. It is a colorless weak organic acid. It occurs naturally in citrus fruits. In biochemistry, it is an intermediate in the citric acid cycle, which occurs in ...
and
malic acid) every 2–3 days in order to aid the mobilization and absorption of phosphate. Many species are fire-adapted (
pyrophyte
Pyrophytes are plants which have adapted to tolerate fire.
Fire acts favourably for some species. "Passive pyrophytes" resist the effects of fire, particularly when it passes over quickly, and hence can out-compete less resistant plants, which are ...
s), meaning they have strategies for surviving fires that sweep through their habitat. Some are
resprouters, and have a thick rootstock buried in the ground that shoots up new stems after a fire, and others are
reseeders, 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.
There are four dioecious genera (''
Aulax
'' Aulax'' is a South African Proteaceae genus of just three species of evergreen shrubs commonly known as "featherbushes". The name ''Aulax'' was named by a botanist named Linnaeus; he named ''Aulax'' after the Greek god Proteus, who is known to ...
'', ''
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
A pollen-presenter is an area on the tip of the style in flowers of plants of the family Proteaceae on which the anthers release their pollen prior to anthesis. To ensure pollination, the style grows during anthesis, sticking out the pollen-presen ...
, 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
Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
is carried out by
bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s,
beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s,
flies,
moths, birds (
honeyeaters,
sunbirds,
sugarbird
The sugarbirds are a small genus, ''Promerops'', and family, Promeropidae, of passerine birds, restricted to southern Africa. In general appearance and habits, they resemble large, long-tailed sunbirds, but are possibly more closely related t ...
s and
hummingbirds) and mammals (rodents, small
marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s,
elephant shrews and
bats
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bir ...
). 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 (
bracts) 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
elaiosomes that are dispersed by
ants; the seeds with wings or thistledown exhibit
anemochory, while the drupes and other fleshy 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
Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
,
India
India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...
, eastern and south eastern
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, and
Oceania
Oceania (, , ) is a geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern and Western hemispheres, Oceania is estimated to have a land area of and a population of around 44.5 million ...
.
Only two species are known from New Zealand, although fossil pollen evidence suggests there were more previously.
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
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. 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
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
coal
Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen.
Coal is formed when ...
deposits of the
South Island of
New Zealand
New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
. It is thought to have achieved its present distribution largely by
continental drift rather than dispersal across ocean gaps.
Phytochemistry
No conclusive studies have been carried out on the chemical substances present in this broad family. The genera ''
Protea'' and ''
Faurea
''Faurea'' is a genus containing 16 species of flowering plants in the protea family which occur in the summer rainfall area of southern Africa, extending to tropical Africa and Madagascar. The name honours South African soldier and botanist
...
'' are unusual as they use
xylose
Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
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
-Tyrosine or tyrosine (symbol Tyr or Y) or 4-hydroxyphenylalanine is one of the 20 standard amino acids that are used by cells to synthesize proteins. It is a non-essential amino acid with a polar side group. The word "tyrosine" is from the G ...
, are often present, as are
proanthocyanidines (
delphinidin and
cyanidin),
flavonols (
kaempferol,
quercetin and
myricetin
Myricetin is a member of the flavonoid class of polyphenolic compounds, with antioxidant properties. Common dietary sources include vegetables (including tomatoes), fruits (including oranges), nuts, berries, tea, and red wine. Myricetin is struc ...
) and
arbutin.
Alkaloid
Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s are usually absent.
Iridoid
Iridoids are a type of monoterpenoids in the general form of cyclopentanopyran, found in a wide variety of plants and some animals. They are biosynthetically derived from 8-oxogeranial. Iridoids are typically found in plants as glycosides, mo ...
s and
ellagic acid
Ellagic acid is a polyphenol found in numerous fruits and vegetables. It is the dilactone of hexahydroxydiphenic acid.
Name
The name comes from the French term ''acide ellagique'', from the word ''galle'' spelled backwards because it can be ...
are also absent.
Saponins and
sapogenin
Sapogenins are the aglycones, or non-saccharide, portions of the family of natural products known as saponins. Sapogenins contain steroid or other triterpene frameworks as their key organic feature. For example, steroidal sapogenins such as tigge ...
s can be either present or absent in different species. Many species accumulate
aluminium
Aluminium (aluminum in American and Canadian English) is a chemical element with the symbol Al and atomic number 13. Aluminium has a density lower than those of other common metals, at approximately one third that of steel. It ha ...
.
Uses and cultivation
Many traditional 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 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 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, 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
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 appropriat ...
are used in gardening, particularly genera of ''
Banksia'', ''
Embothrium'', ''
Grevillea'', and ''
Telopea''. Unfortunately, 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
Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
and
New Zealand
New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
, 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. It is also planted in the
British Isles
The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isl ...
and on the Pacific coast of the
United States
The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
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 flowers and foliage. Some species are of importance to the
cut flower industry
Cut flowers are flowers or flower buds (often with some stem and leaf) that have been cut from the plant bearing it. It is usually removed from the plant for decorative use. Typical uses are in vase displays, wreaths and garlands. Many ga ...
, 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
'' Aulax'' is a South African Proteaceae genus of just three species of evergreen shrubs commonly known as "featherbushes". The name ''Aulax'' was named by a botanist named Linnaeus; he named ''Aulax'' after the Greek god Proteus, who is known to ...
'' and blushing brides (''
Serruria
''Serruria'', or spiderhead is a genus of flowering plants in the family Proteaceae, endemic to South Africa.
Names
''Serruria'' was named in honor of Joseph Serrurier, a professor of botany at the Utrecht University early in the eighteent ...
''), 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
A wildlife garden (or wild garden) is an environment created by a gardener that serves as a sustainable haven for surrounding wildlife. Wildlife gardens contain a variety of habitats that cater to native and local plants, birds, amphibians, ...
s 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
In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
a tropical appearance; ''
Lomatia ferruginea'' (Fuinque), ''
Lomatia hirsuta
Lomatia hirsuta, known as radal, is a tree native to Argentina, Chile, and Peru.
In Chile it grows from Coquimbo to Chiloe (36 to 42°S). It grows on a variety of soils and humidity. It grows on poor soils near ravines.
Description
''Lomatia ...
'' (Radal) have been introduced in Western
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and to the western
United States
The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
. ''
Embothrium coccineum
''Embothrium coccineum'', Chilean firetree or Chilean firebush, commonly known in Chile and Argentina as ''notro'', ''ciruelillo'' and ''fósforo'' is a small evergreen tree in the flowering plant family Proteaceae. It grows in the temperate for ...
'' (Chilean Firetree or ''Notro'') is highly valued in the British Isles for its dark red flowers and can be found as far north as the
Faroe Islands
The Faroe Islands ( ), or simply the Faroes ( fo, Føroyar ; da, Færøerne ), are a North Atlantic island group and an autonomous territory of the Kingdom of Denmark.
They are located north-northwest of Scotland, and about halfway bet ...
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 flowers.
Another species that is cultivated in some parts of the world, although it is smaller, is ''
Telopea speciosissima
''Telopea speciosissima'', commonly known as the New South Wales waratah or simply waratah, is a large shrub in the plant family Proteaceae. It is endemic to New South Wales in Australia and is the floral emblem of that state. No subspecies ...
'' (Waratah), from the mountains of
New South Wales
)
, nickname =
, image_map = New South Wales in Australia.svg
, map_caption = Location of New South Wales in AustraliaCoordinates:
, subdivision_type = Country
, subdivision_name = Australia
, established_title = Before federation
, es ...
,
Australia
Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
.
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 flowers and grape-like fruit. ''
Adenanthos sericeus
''Adenanthos sericeus'', commonly known as woolly bush, is a shrub native to the south coast of Western Australia. It has bright red but small and obscure flowers, and very soft, deeply divided, hairy leaves.
Description
''Adenanthos sericeus' ...
'' (woolly bush) is planted for its attractive soft leaves and its small red or orange flowers. ''
Hicksbeachia pinnatifolia'' (beef nut, red bauple nut) is commonly planted for its foliage and edible nuts.
Parasites
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
''Calonectria'' (anamorph '' Cylindrocladium'') is a genus of ascomycete fungi. ''Calonectria'' species are plant pathogens
Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectio ...
'' and other fungi.
Conservation status
The
IUCN[IUCN 2006. 2006 IUCN Red List of Threatened Species. . Downloaded on 22 February 2007] 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
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
, fire, parasitic diseases,
competition
Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, ind ...
from introduced plants,
soil degradation and other damage provoked by humans and their domesticated animals. The species are also affected by
climate change.
Fossils
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
Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a p ...
bases or, in other cases, the unusual structure of pollen tetrads. ''
Xylocaryon
''Xylocaryon'' is an extinct genus of plants in the family Proteaceae. The sole species is ''Xylocaryon lockii'' from south-eastern Australia, described from fossilised fruits found at Nintingbool near Ballarat, Victoria and Flinders Island
...
'' was identified as a member of the Proteaceae from the similarity of its fruit to the extant genus ''
Eidothea
''Eidothea'' is a genus of two species of rainforest trees in New South Wales and Queensland, in eastern Australia, constituting part of the plant family Proteaceae. The plant family Proteaceae was named after the shape-shifting god Proteus of G ...
''.
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
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
(
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.
Taxonomy
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
taxonomists
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
. 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.
A classification of the genera within Proteaceae was made by
Lawrie Johnson and
Barbara 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 ...
in their influential 1975 monograph "
On the Proteaceae: the evolution and classification of a southern family",
until it was largely superseded by the molecular studies of Peter H. Weston and Nigel Barker in 2006. Proteaceae are now divided into five subfamilies:
Bellendenoideae,
Persoonioideae,
Symphionematoideae,
Proteoideae and
Grevilleoideae
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 ...
.
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''.
The full arrangement, according to Weston and Barker (2006) with the updates to genera from Mast et al. (2008), is as follows:
:Family Proteaceae
::Subfamily
Bellendenoideae
:::::''
Bellendena
''Bellendena montana'', commonly known as mountain rocket, is a species of low-growing multi-stemmed shrub in the plant family Proteaceae. It is endemic to high-altitude subalpine and alpine regions in Tasmania, Australia. The prominent white fl ...
''
::Subfamily
Persoonioideae
:::Tribe
Placospermeae
:::::''
Placospermum''
:::Tribe
Persoonieae
:::::''
Toronia
''Toronia'' is a genus of tree in the family Proteaceae that contains a single species, ''Toronia toru'', which is endemic to New Zealand. The genus is closely related to the large genus ''Persoonia'', and in fact this species was long regarded ...
'' — ''
Garnieria
''Garnieria'' is a monotypic plant genus in the family Proteaceae. Molecular hylogenetic studies indicate that it is nested in the larger Australian genus ''Persoonia'' where it may be included. It is endemic to New Caledonia where the type speci ...
'' — ''
Acidonia
''Acidonia microcarpa'' is a species of shrub in the plant family Proteaceae. It is the only species in the genus ''Acidonia''. It is endemic to the south coast of the Southwest Botanic Province of Western Australia.
It was originally publishe ...
'' — ''
Persoonia''
::Subfamily
Symphionematoideae
:::::''
Agastachys
''Agastachys odorata'', commonly known as the white waratah or fragrant candlebush, is the sole member of the genus ''Agastachys'' in the protea family. It is an evergreen shrub to small tree and is endemic to the heaths and button grass sedgela ...
'' — ''
Symphionema''
::Subfamily
Proteoideae
:::
incertae sedis
:::::''
Eidothea
''Eidothea'' is a genus of two species of rainforest trees in New South Wales and Queensland, in eastern Australia, constituting part of the plant family Proteaceae. The plant family Proteaceae was named after the shape-shifting god Proteus of G ...
'' — ''
Beauprea'' — ''
Beaupreopsis'' — ''
Dilobeia'' — ''
Cenarrhenes'' — ''
Franklandia''
:::Tribe
Conospermeae
::::Subtribe
Stirlingiinae
:::::''
Stirlingia''
::::Subtribe
Conosperminae
:::::''
Conospermum'' — ''
Synaphea
''Synaphea'' is a genus of small shrubs and is Endemism, endemic to Western Australia. Synapheas have variably shaped leaves but consistently yellow flowers with an unusual pollination mechanism.
Description
Plants in the genus ''Synaphea'' are ...
''
:::Tribe
Petrophileae
:::::''
Petrophile
''Petrophile'' is a genus of evergreen shrubs, in the family Proteaceae. The genus is endemic to Australia. Commonly known as conebushes, they typically have prickly, divided foliage and produce prominently-displayed pink, yellow or cream flow ...
'' — ''
Aulax
'' Aulax'' is a South African Proteaceae genus of just three species of evergreen shrubs commonly known as "featherbushes". The name ''Aulax'' was named by a botanist named Linnaeus; he named ''Aulax'' after the Greek god Proteus, who is known to ...
''
:::Tribe
Proteeae
:::::''
Protea'' — ''
Faurea
''Faurea'' is a genus containing 16 species of flowering plants in the protea family which occur in the summer rainfall area of southern Africa, extending to tropical Africa and Madagascar. The name honours South African soldier and botanist
...
''
:::Tribe
Leucadendreae
::::Subtribe
Isopogoninae
:::::''
Isopogon
''Isopogon'', commonly known as conesticks, conebushes or coneflowers, is a genus of about forty species of flowering plants in the family Proteaceae, and are endemic to Australia. They are shrubs with rigid leaves, bisexual flowers in a dense ...
''
::::Subtribe
Adenanthinae
:::::''
Adenanthos
''Adenanthos'' is a genus of Australian native shrubs in the flowering plant family Proteaceae. Variable in habit and leaf shape, it is the only genus in the family where solitary flowers are the norm. It was discovered in 1791, and formally publ ...
''
::::Subtribe
Leucadendrinae
:::::''
Leucadendron'' — ''
Serruria
''Serruria'', or spiderhead is a genus of flowering plants in the family Proteaceae, endemic to South Africa.
Names
''Serruria'' was named in honor of Joseph Serrurier, a professor of botany at the Utrecht University early in the eighteent ...
'' — ''
Paranomus'' — ''
Vexatorella'' — ''
Sorocephalus'' — ''
Spatalla
''Spatalla'' is a genus containing 20 species of flowering plants, commonly known as "spoons", in the family Proteaceae. The genus is endemic to the Cape Floristic Region of South Africa where it is associated with fynbos habitats. The spe ...
'' — ''
Leucospermum'' — ''
Mimetes'' — ''
Diastella'' — ''
Orothamnus''
::Subfamily
Grevilleoideae
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 ...
:::
incertae sedis
:::::''
Sphalmium'' — ''
Carnarvonia''
:::Tribe
Roupaleae
::::
incertae sedis
:::::''
Megahertzia'' — ''
Knightia'' — ''
Eucarpha'' — ''
Triunia''
::::Subtribe
Roupalinae
:::::''
Roupala'' — ''
Neorites'' — ''
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 o ...
''
::::Subtribe
Lambertiinae
:::::''
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 s ...
'' — ''
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 pair ...
''
::::Subtribe
Heliciinae
:::::''
Helicia'' — ''
Hollandaea''
::::Subtribe
Floydiinae
:::::''
Darlingia'' — ''
Floydia''
:::Tribe
Banksieae
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 ...
::::Subtribe
Musgraveinae
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
Th ...
:::::''
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 relat ...
'' — ''
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, Austral ...
''
::::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 i ...
:::::''
Banksia''
:::Tribe
Embothrieae
::::Subtribe
Lomatiinae
:::::''
Lomatia''
::::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
The ...
:::::''
Embothrium'' — ''
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
:::::''
Stenocarpus'' — ''
Strangea''
::::Subtribe
Hakeinae
:::::''
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. celsis ...
'' — ''
Hakea'' — ''
Grevillea'' — ''
Finschia''
:::Tribe
Macadamieae
::::Subtribe
Macadamiinae
:::::''
Macadamia'' — ''
Lasjia'' — ''
Nothorites'' — ''
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 magnifr ...
'' — ''
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
:::::''
Malagasia'' — ''
Catalepidia''
::::Subtribe
Virotiinae
:::::''
Virotia
''Virotia'' is a genus of six species of flowering plants in the family Proteaceae. The genus is endemic to New Caledonia with six species that were once placed in ''Macadamia''.Virot, R. (1967). Protéacées. In Flore de La Nouvelle-Calédonie e ...
'' — ''
Athertonia'' — ''
Heliciopsis''
::::Subtribe
Gevuininae
:::::''
Cardwellia'' — ''
Sleumerodendron'' — ''
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". Plan ...
'' — ''
Gevuina'' — ''
Bleasdalea'' — ''
Hicksbeachia'' — ''
Kermadecia'' — ''
Turrillia''
References
*
*
*
*
* Brown, R
On the Proteaceae of Jussieu''
Proceedings of the Linnean Society'' 10:15-216.
External links
*
*
Images of Proteaceae from the Australian National Botanical GardensMap
{{Authority control
Eudicot families
Extant Cenomanian first appearances