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''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen
coniferous Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extant ...
trees in the family Araucariaceae. There are 20 
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
species in
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
(where 14 species are endemic, see New Caledonian ''Araucaria''),
Norfolk Island Norfolk Island (, ; Norfuk: ''Norf'k Ailen'') is an external territory of Australia located in the Pacific Ocean between New Zealand and New Caledonia, directly east of Australia's Evans Head and about from Lord Howe Island. Together with ...
, eastern
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, New Guinea, East Argentina, South Brazil, Chile and Paraguay. They are still common in the
South Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
rejoin and Eastern Australia.


Description

''Araucaria'' are mainly large trees with a massive erect stem, reaching a height of . The horizontal, spreading branches grow in whorls and are covered with leathery or needle-like
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
. In some species, the leaves are narrow, awl-shaped and lanceolate, barely overlapping each other; in others they are broad and flat, and overlap broadly. The trees are mostly
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
, with male and female cones found on separate trees, though occasional individuals are
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
or change sex with time. The female cones, usually high on the top of the tree, are
globose A sphere () is a geometrical object that is a three-dimensional analogue to a two-dimensional circle. A sphere is the set of points that are all at the same distance from a given point in three-dimensional space.. That given point is the ce ...
, and vary in size among species from diameter. They contain 80–200 large edible seeds, similar to pine nuts, though larger. The male cones are smaller, long, and narrow to broad cylindrical, broad. The genus is familiar to many people as the genus of the distinctive Chilean pine or monkey-puzzle tree ('' Araucaria araucana''). The genus is named after the Spanish exonym ''Araucano'' ("from Arauco") applied to the Mapuche of south-central Chile and south-west Argentina, whose territory incorporates natural stands of this genus. The Mapuche people call it ', and consider it sacred. Some Mapuche living in the Andes name themselves Pehuenche ("people of the '") as they traditionally harvested the seeds extensively for food. No distinct vernacular name exists for the genus. Many are called "pine", although they are only distantly related to true pines, in the genus '' Pinus''.


Distribution and paleoecology

Members of ''Araucaria'' are found in Argentina, Brazil,
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
,
Norfolk Island Norfolk Island (, ; Norfuk: ''Norf'k Ailen'') is an external territory of Australia located in the Pacific Ocean between New Zealand and New Caledonia, directly east of Australia's Evans Head and about from Lord Howe Island. Together with ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, New Guinea, Chile and
Papua (Indonesia) Papua is a province of Indonesia, comprising the northern coast of Western New Guinea together with island groups in Cenderawasih Bay to the west. It roughly follows the borders of Papuan customary region of Tabi Saireri. It is bordered by the ...
. Many if not all current populations are relicts, and of restricted distribution. They are found in forest and
maquis shrubland 220px, Low maquis in Corsica 220px, High ''macchia'' in Sardinia ( , , ) or ( , ; often in Italian; hr, makija; ; ) is a shrubland biome in the Mediterranean region, typically consisting of densely growing evergreen shrubs. Maquis is char ...
, with an affinity for exposed sites. The earliest records of the genus date to the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
, represented by ''
Araucaria mirabilis ''Araucaria mirabilis'' is an extinct species of coniferous tree from Patagonia, Argentina. It belongs to the genus '' Araucaria''. ''A. mirabilis'' are known from large amounts of very well preserved silicified wood and cones from the Cerro ...
'' of Argentina. Fossil records show that the genus also formerly occurred in the northern hemisphere until the end of the Cretaceous period. By far the greatest diversity exists in
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
, likely due to a relatively recent
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 int ...
, as all New Caledonian species are more closely related to each other than they are to other ''Araucaria''. Much of New Caledonia is composed of ultramafic rock with
serpentine soil Serpentine soil is an uncommon soil type produced by weathered ultramafic rock such as peridotite and its metamorphic derivatives such as serpentinite. More precisely, serpentine soil contains minerals of the serpentine subgroup, especially anti ...
s, with low levels of nutrients, but high levels of metals such as nickel. Consequently, its endemic ''Araucaria'' species are adapted to these conditions, and many species have been severely affected by
nickel mining in New Caledonia Nickel mining in New Caledonia is a major sector of the New Caledonian economy. The island contains about 7.1 million tonnes of nickel reserves, about 10% of the world's total. With an annual production of 200,000 tonnes in 2020, New Caledonia wa ...
and are now considered threatened or endangered, due to their habitat lying in prime areas for nickel mining activities. Some evidence suggests that the long necks of
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
dinosaurs may have evolved specifically to browse the foliage of tall trees, including those of ''Araucaria''. An analysis of modern ''Araucaria'' leaves found that they have a high energy content but are slow fermenting, making their ancestors a likely attractive target.


Classification and species list

There are four extant
sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
and two extinct
sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
in the genus, sometimes treated as separate genera.


Extant Species


Genetic studies

Genetic studies indicate that the extant members of the genus can be subdivided into two large
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s – the first consisting of the
sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
''Araucaria'', ''Bunya'', and ''Intermedia''; and the second of the strongly
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
section ''Eutacta''. Sections ''Eutacta'' and ''Bunya'' are both the oldest taxa of the genus, with ''Eutacta'' possibly older. ::Taxa marked with are extinct. * Section ''Araucaria''. Leaves broad; cones more than diameter; seed germination hypogeal. Syn. sect. ''Columbea''; sometimes includes ''Intermedia'' and ''Bunya'' ** ''
Araucaria angustifolia ''Araucaria angustifolia'', the Paraná pine, Brazilian pine or candelabra tree (, or ), is a critically endangered species in the conifer genus '' Araucaria''. Although the common names in various languages refer to the species as a "pine", it d ...
'' – Paraná pine (obsolete: Brazilian pine, candelabra tree); southern and southeastern Brazil, northeastern Argentina. ** '' Araucaria araucana'' – monkey-puzzle or ''pehuén'' (obsolete: Chile pine); central Chile & western Argentina. ** ''
Araucaria nipponensis ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20  extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Arauca ...
'' – Japan and Sakhalin (Upper Cretaceous) * Section ''Bunya''. Contains only one living species. Produces recalcitrant seeds with hypogeal ( cryptocotylar) germination, though extinct species may have exhibited epigeal germination. ** '' Araucaria bidwillii'' – ''bunya-bunya''; Eastern Australia * Section ''Intermedia''. Contains only one living species. Produces recalcitrant seeds ** '' Araucaria hunsteinii'' – ''klinki''; New Guinea ** ''
Araucaria haastii ''Araucaria haastii'' is an extinct species of conifer tree formerly native to New Zealand. A large number of fossilised tree specimens from the family Araucariaceae have been found in New Zealand, but in many cases the level of preservation is ...
'' - New Zealand (Cretaceous) * Section ''Eutacta''. Leaves narrow, awl-like; cones less than diameter; seed germination
epigeal Epigeal, epigean, epigeic and epigeous are biological terms describing an organism's activity above the soil surface. In botany, a seed is described as showing epigeal germination when the cotyledons of the germinating seed expand, throw off the ...
** '' Araucaria bernieri'' – New Caledonia ** ''
Araucaria biramulata ''Araucaria biramulata'', the biramule araucaria, or piggyback araucaria, is a species of conifer in the family Araucariaceae. It is found only in New Caledonia on the main island of Grande Terre (New Caledonia), Grande Terre. ''Araucaria biramu ...
'' – New Caledonia ** '' Araucaria columnaris'' – Cook pine; New Caledonia ** '' Araucaria cunninghamii'' – Moreton Bay pine, hoop pine; Eastern Australia, New Guinea ** ''
Araucaria goroensis ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Araucaria ...
'' – New Caledonia ** '' Araucaria heterophylla'' – Norfolk Island pine; Norfolk Island ** '' Araucaria humboldtensis'' – New Caledonia ** '' Araucaria laubenfelsii'' – New Caledonia ** ''
Araucaria luxurians ''Araucaria luxurians'' is a species of conifer in the family Araucariaceae. It is known by the common names Sapin de Noël and Coast araucaria.
'' – New Caledonia ** ''
Araucaria montana ''Araucaria montana'', the mountain araucaria, is a species of conifer in the family Araucariaceae. It is found only in New Caledonia. ''Araucaria montana'' is the most widely distributed of New Caledonia's native ''Araucaria'' species, and is f ...
'' – New Caledonia ** ''
Araucaria muelleri ''Araucaria muelleri'' is a species of conifer in the family Araucariaceae. It is a medium size tree, 10–25 meters in height, with larger leaves than most other New Caledonian ''Araucarias''. It is found only in New Caledonia, in several sites i ...
'' – New Caledonia ** '' Araucaria nemorosa'' – New Caledonia ** ''
Araucaria rulei ''Araucaria rulei'' (Rule araucaria, graceful pine) is a species of conifer in the family Araucariaceae. It is endemic to New Caledonia, where it is an endangered species. Its populations are fragmented and are generally made up of scattered ...
'' – New Caledonia ** '' Araucaria schmidii'' – New Caledonia ** ''
Araucaria scopulorum ''Araucaria scopulorum'' (rock araucaria) is a species of conifer in the family Araucariaceae. It is endemic to New Caledonia, where it is an endangered species. It occurs in small populations that are restricted to two main areas. It grows in sh ...
'' – New Caledonia ** ''
Araucaria subulata ''Araucaria subulata'' (narrow-leaf araucaria) is a species of conifer in the family Araucariaceae. It is found only in New Caledonia, with scattered populations present across the central and southern mountain regions of the main island Grand ...
'' – New Caledonia ** '' Araucaria lignitici'' – (Paleogene) Yallourn, Victoria, Australia **'' Araucaria famii –'' (Late Cretaceous) Vancouver Island, Canada. * Section ''Yezonia''. Extinct. Contains only one species ** ''
Araucaria vulgaris ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Ara ...
'' – Japan (Late Cretaceous) * Section ''Perpendicula''. Extinct. Contains only one species ** '' Araucaria desmondii'' - New Zealand (Late Cretaceous) * incertae sedis ** ''
Araucaria beipiaoensis ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20  extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Arauca ...
'' – Tiaojishan Formation, China (Middle Jurassic) ** ''
Araucaria fibrosa ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen Conifer, coniferous trees in the family Araucariaceae. There are 20 extant taxon, extant species in New Caledonia (where 14 species are endemism, ende ...
'' –
López de Bertodano Formation The Lopez de Bertodano Formation is a geological Formation (geology), formation in the James Ross Island, James Ross archipelago of the Antarctic Peninsula. The strata date from the end of the Late Cretaceous (upper-lower Maastrichtian stage) to t ...
, Antarctica (Late Cretaceous) ** '' Araucaria marensii'' –
La Meseta Formation The La Meseta Formation is a sedimentary sequence deposited during the Eocene. The formation is found on Seymour Island, Antarctica. Description La Meseta Formation lies unconformably on the Cretaceous Lopez de Bertodano Formation. It is an app ...
, Antarctica & Santa Cruz Formation, Argentina''Araucaria marensii''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org
** ''
Araucaria nihongii ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20  extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Arauca ...
'' – Upper
Yezo Group The Yezo Group is a stratigraphic group in Hokkaido, Japan and Sakhalin, Russia which is primarily Late Cretaceous in age (Aptian to Earliest Paleocene). It is exposed as roughly north–south trending belt extending 1,500 kilometres through centr ...
, Japan (Late Cretaceous) ** ''
Araucaria taieriensis ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Araucari ...
'' - New Zealand (Late Cretaceous) **''
Araucaria brownii ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Ara ...
'' - England (Middle Jurassic) ** ''
Araucaria mirabilis ''Araucaria mirabilis'' is an extinct species of coniferous tree from Patagonia, Argentina. It belongs to the genus '' Araucaria''. ''A. mirabilis'' are known from large amounts of very well preserved silicified wood and cones from the Cerro ...
'' – Patagonia (Middle Jurassic) ** ''
Araucaria sphaerocarpa ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Araucaria ...
'' - England (Middle Jurassic) ''Araucaria bindrabunensis'' (previously classified under section ''Bunya'') has been transferred to the genus '' Araucarites''.


Uses

Some of the species are relatively common in cultivation because of their distinctive, formal symmetrical growth habit. Several species are economically important for timber production.


Food

The edible large seeds of ''A. araucana'', ''A. angustifolia'' and ''A. bidwillii'' — also known as ''Araucaria'' nuts, and often called, although improperly, '' pine nuts'' — are eaten as food, particularly among the Mapuche people of Chile and southwest Argentina and among
Native Australian Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples ...
s. In South America ''Araucaria'' nuts or seeds are called ''piñas'', ''pinhas'', ''piñones'' or ''pinhões'', like pine nuts in Europe.


Pharmacological activity

Pharmacological reports on genus Araucaria are anti-ulcer, antiviral, neuro-protective, anti-depressant and
anti-coagulant Anticoagulants, commonly known as blood thinners, are chemical substances that prevent or reduce coagulation of blood, prolonging the clotting time. Some of them occur naturally in blood-eating animals such as leeches and mosquitoes, where they ...
.


See also

* ''
Agathis ''Agathis'', commonly known as kauri or dammara, is a genus of 22 species of evergreen tree. The genus is part of the ancient conifer family Araucariaceae, a group once widespread during the Jurassic and Cretaceous periods, but now largely re ...
'' (kauri) * '' Wollemia''


References


External links

* * * * {{Taxonbar, from=Q157214 Dioecious plants Extant Triassic first appearances Conifer genera