Araucaria
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Araucaria (Template:IPAc-en; original pronunciation: [a.ɾawˈka. ɾja])<ref>Template:OED</ref> is a genus of evergreen coniferous trees in the family Araucariaceae. While today they are largely confined to the Southern Hemisphere, during the Jurassic and Cretaceous they were globally distributed. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian Araucaria), eastern Australia (including Norfolk Island), New Guinea, Argentina, Brazil, Chile and Uruguay.
The genus is familiar to many people as the genus of the distinctive Chilean pine or monkey-puzzle tree (Araucaria araucana). 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.
Description
Araucaria are mainly large trees with a massive erect stem, reaching a height of Template:Convert. The horizontal, spreading branches grow in whorls and are covered with leathery or needle-like leaves. 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.<ref name="gymdb"/>
The trees are mostly dioecious, with male and female cones found on separate trees,<ref name="pinetum">Template:Cite web</ref> though occasional individuals are monoecious or change sex with time.<ref name="simmons">Template:Cite book</ref> The female cones, usually high on the top of the tree, are globose, and vary in size among species from Template:Convert in diameter. They contain 80–200 large edible seeds, similar to pine nuts, though larger. The male cones are smaller, Template:Convert long, narrow to broad cylindrical, and Template:Convert broad.
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Taxonomy
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 a species in this genus identified as A. araucana; the Mapuche people call it Template:Lang, and consider it sacred.<ref name="gymdb"/> Some Mapuche living in the Andes name themselves Pewenche ("people of the Template:Lang") as they traditionally harvested the seeds extensively for food.<ref name="ntbg">Template:Cite web</ref><ref name="moreno">Template:Cite web</ref>
There are four extant sections and two extinct sections in the genus, sometimes treated as separate genera.<ref name="gymdb"/><ref name="black">Template:Cite book</ref><ref name="Eckenwalder">Template:Cite book</ref>
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Extant species
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Genetic studies
Genetic studies indicate that the extant members of the genus can be subdivided into two large clades – the first consisting of the sections Araucaria, Bunya, and Intermedia; and the second of the strongly monophyletic section Eutacta. Sections Eutacta and Bunya are both the oldest taxa of the genus, with Eutacta possibly older.<ref name="setoguchi">Template:Cite journal</ref>
- Taxa marked with Template:Extinct are extinct.
- Template:Anchor Section Araucaria. Template:Small Leaves broad; cones more than Template:Convert diameter; seed germination hypogeal. Syn. sect. Columbea; sometimes includes Intermedia and Bunya
- Araucaria angustifolia – 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.
- Template:ExtinctAraucaria nipponensis – Japan and Sakhalin (Upper Cretaceous)<ref name="dettman">Template:Cite book</ref>
- Template:Anchor Section Bunya. Template:Small Contains only one living species. Produces recalcitrant seeds with hypogeal (cryptocotylar) germination,<ref name="gotz">Template:Cite book</ref> though extinct species may have exhibited epigeal germination.<ref name="setoguchi"/>
- Araucaria bidwillii – bunya-bunya; Eastern Australia
- Template:Anchor Section Intermedia. Template:Small Contains only one living species. Produces recalcitrant seeds
- Araucaria hunsteinii – klinki; New Guinea
- Template:ExtinctAraucaria haastii - New Zealand (Cretaceous)
- Template:Anchor Section Eutacta. Template:Small Leaves narrow, awl-like; cones less than Template:Convert diameter; seed germination epigeal
- Araucaria bernieri – New Caledonia
- Araucaria biramulata – New Caledonia
- Araucaria columnaris – Cook pine; New Caledonia
- Araucaria cunninghamii – Moreton Bay pine, hoop pine; Eastern Australia, New Guinea
- Araucaria goroensis – New Caledonia
- Araucaria heterophylla – Norfolk Island pine; Norfolk Island
- Araucaria humboldtensis – New Caledonia
- Araucaria laubenfelsii – New Caledonia
- Araucaria luxurians – New Caledonia
- Araucaria montana – New Caledonia
- Araucaria muelleri – New Caledonia
- Araucaria nemorosa – New Caledonia
- Araucaria rulei – New Caledonia
- Araucaria schmidii – New Caledonia
- Araucaria scopulorum – New Caledonia
- Araucaria subulata – New Caledonia
- Template:ExtinctAraucaria lignitici – (Paleogene) Yallourn, Victoria, Australia<ref name="Cookson 1951">Template:Cite journal</ref>
- Template:ExtinctAraucaria famii – (Late Cretaceous) Vancouver Island, Canada.<ref name=":0">Template:Cite journal</ref>
- Template:ExtinctSection Yezonia. Extinct. Contains only one species
- Template:ExtinctAraucaria vulgaris – Japan (Late Cretaceous)
- Template:ExtinctSection Perpendicula. Extinct. Contains only one species
- Template:ExtinctAraucaria desmondii - New Zealand (Late Cretaceous)
- incertae sedis
- Template:ExtinctAraucaria antarctica - Snow Hill Island Formation, Antarctica (Late Cretaceous)<ref>Template:Cite journal</ref>
- Template:ExtinctAraucaria beipiaoensis – Tiaojishan Formation, China (Middle Jurassic)
- Template:ExtinctAraucaria fibrosa – López de Bertodano Formation, Antarctica (Late Cretaceous)
- Template:ExtinctAraucaria marensii – La Meseta Formation, Antarctica & Santa Cruz Formation, Argentina<ref name=FWAraucariaMarensii>Araucaria marensii at Fossilworks.org</ref><ref name=Vicaino2012_p112>Template:Cite book</ref>
- Template:ExtinctAraucaria nihongii – Upper Yezo Group, Japan (Late Cretaceous)
- Template:ExtinctAraucaria taieriensis - New Zealand (Late Cretaceous)<ref>Template:Cite journal</ref>
- Template:ExtinctAraucaria brownii - England (Middle Jurassic)
- Template:ExtinctAraucaria mirabilis – Patagonia (Middle Jurassic)
- Template:ExtinctAraucaria sphaerocarpa - England (Middle Jurassic)
- Template:ExtinctAraucaria grandifolia - Argentina (Early Cretaceous)<ref>Template:Cite journal</ref>
- Template:ExtinctAraucaria jeffreyi - Eastern United States (Late Cretaceous)<ref>Template:Cite web</ref>
Araucaria bindrabunensis (previously classified under section Bunya) has been transferred to the genus Araucarites.
Distribution and paleoecology
Members of Araucaria are found in Argentina, Brazil, New Caledonia, Norfolk Island, Australia, New Guinea, Chile and Papua (Indonesia).<ref>The Pine Trees of Lanai</ref> Many if not all current populations are relicts, and of restricted distribution. They are found in forest and maquis shrubland, with an affinity for exposed sites. The earliest records of the genus date to the Middle Jurassic, represented by Araucaria mirabilis of Argentina, and Araucaria sphaerocarpa from England. Fossil records show that the genus also formerly occurred in the northern hemisphere until the end of the Cretaceous period.<ref name=":0" /> A possible species of Araucaria may have existed in New Zealand as recently as 350,000 years ago.<ref name=":1">Template:Cite journal</ref> However, it is also possible that this fossil actually represents a specimen of Agathis.<ref name=":1" />
By far the greatest diversity exists in New Caledonia, likely due to a relatively recent adaptive radiation, as all New Caledonian species are more closely related to each other than they are to other Araucaria.<ref name=":0" /><ref name="gymdb">Template:Cite web</ref> Much of New Caledonia is composed of ultramafic rock with serpentine soils, with low levels of nutrients, but high levels of metals such as nickel.<ref>Template:Cite web</ref> Consequently, its endemic Araucaria species are adapted to these conditions, and many species have been severely affected by nickel mining in New Caledonia 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 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.<ref name="hummel">Template:Cite journal</ref>
Uses
Template:Further 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.
FoodTemplate:Anchor
Template:Broader The edible large seeds of A. araucana, A. angustifolia and A. bidwillii — also known as Araucaria nuts,<ref>Template:Cite encyclopedia</ref> and often called, although improperly, pine nuts — are eaten as food, particularly among the Mapuche people of Chile and southwest Argentina, the Kaingang people in Southern Brazil and among Indigenous Australians.<ref name="gymdb"/> In South America Araucaria nuts or seeds are called piñas or piñones in Spanish and pinhões in Portuguese, like pine nuts in Europe.
Pharmacological activity
Pharmacological reports on the genus Araucaria are anti-ulcer, antiviral, neuro-protective, anti-depressant and anti-coagulant.<ref>Template:Cite journal</ref>
See also
References
External links
Template:Nuts Template:Plant classification Template:Acrogymnospermae classification Template:Taxonbar Template:Authority control