Araucariaceae
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Araucariaceae is a family of conifers with three living genera, Araucaria, Agathis, and Wollemia. While the family's native distribution is now largely confined to the Southern Hemisphere, except for a few species of Agathis in Malesia, it was formerly widespread in the Northern Hemisphere during the Jurassic and Cretaceous periods.<ref>Template:Cite journal</ref>
Description
Members of Araucariaceae are typically extremely tall evergreen trees,<ref name="gym">Template:Cite web</ref> reaching heights of Template:Convert or more.<ref name="ucmp">Template:Cite web</ref> They can also grow very large stem diameters; a New Zealand kauri tree (Agathis australis) named Tāne Mahuta ("The Lord of the Forest") has been measured at Template:Convert tall with a diameter at breast height of Template:Convert. Its total wood volume is calculated to be Template:Convert,<ref name="kauri">Template:Cite web</ref> making it the third-largest conifer after Sequoia and Sequoiadendron (both from the Cupressaceae subfamily Sequoioideae).<ref name="gym"/>
The trunks are columnar and have relatively large piths with resinous cortices.<ref name="wu">Template:Cite journal</ref> The branching is usually horizontal and tiered, arising regularly in whorls of three to seven branches or alternating in widely separated pairs.<ref name="ecken">Template:Cite book</ref>
The leaves can be small, needle-like, and curved, or they can be large, broadly ovate, and flattened.<ref name="walters">Template:Cite book</ref> They are spirally arranged, persistent, and usually have parallel venation.<ref name="gym"/>
Like other conifers, they produce cones. Each tree can have both male and female cones (monoecious) or, more commonly, they can have only male or female cones (dioecious).<ref name="carr">Template:Cite web</ref>
Male cones are among the largest among all conifer cones, on average. They are cylindrical and drooping, somewhat resembling catkins. They are borne singly on the tips of branches or in the axils of leaves. They contain numerous sporophylls arranged in whorls or spirals. Each has four to 20 elongated pollen sacs attached to the lower surface at one end. The pollen grains are round and do not possess wings or air sacs.<ref name="gym"/><ref name="ecken"/><ref name="walters"/>
Female cones are also very large. They are spherical to ovoid in shape and borne erect on thick, short shoots at branch tips. The numerous bracts and scales are either fused to each other or separate for half of their lengths.<ref name="gym"/><ref name="ecken"/><ref name="walters"/> The scales almost always bear only one seed on its upper surface, in contrast to two in true pines (family Pinaceae).<ref name="ww">Template:Cite web</ref> They are very large, among the largest seeds among conifers. They are dispersed by wind, usually using wing-like structures. On maturity, the female cones detach and fall to the ground.<ref name="gym"/><ref name="ecken"/><ref name="walters"/> Due to their size, they can cause serious injuries if they hit a person. The cones of the bunya bunya, Araucaria bidwillii, for example, weigh up to Template:Cvt,<ref name="TGD">Template:Cite web</ref> about the size and weight of a large pineapple. They can drop from heights of Template:Convert.<ref name="ww"/>
Classification and genera
Araucariaceae is classified under the order Pinales, class Pinopsida of the division Pinophyta. The division includes all living conifers. Recently however, some authorities treat Araucariaceae as a separate order, Araucariales.<ref name="gym"/>
Araucariaceae contains three extant genera and about 41 species.<ref name="wu"/>
| Image | Genus | Living Species | Distribution |
|---|---|---|---|
|
Araucaria Jussieu |
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19 living species found in New Caledonia (where 13 species are endemic), Norfolk Island, Australia, New Guinea, Argentina, Chile, and Brazil. |
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Agathis Salisbury |
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New Zealand, Australia, Vanuatu, New Caledonia, Papua New Guinea, Indonesia, Malaysia, and the Philippines |
|
Wollemia W.G. Jones, K.D. Hill & J.M. Allen | Endemic to Australia. It was known only from fossil remains before the discovery of the living species in 1994. |
Phylogeny
Below is the phylogeny of the Pinophyta based on cladistic analysis of molecular data. It shows the position of Araucariaceae within the division.<ref>Derived from papers by A. Farjon and C. J. Quinn & R. A. Price in the Proceedings of the Fourth International Conifer Conference, Acta Horticulturae 2003; 615</ref> Template:Clade Relationships between living members of Araucariaceae.<ref name="Escapa 1153–1170">Template:Cite journal</ref>
Template:Clade Molecular evidence supports Araucariaceae and Podocarpaceae having diverged from each other during the late Permian.<ref>Template:Cite journal</ref>
Distribution and habitat
Today, 41 species are known, in three genera: Agathis, Araucaria and Wollemia, distributed largely in the Southern Hemisphere.
By far the greatest diversity is in New Caledonia (18 species), with others in Australia, Argentina, New Zealand, Chile, southern Brazil, and Malesia. In Malesia, Agathis extends a short distance into the Northern Hemisphere, reaching 18°N in the Philippines.
Uses
Template:Further Several species are very popular ornamental trees in gardens in subtropical regions, and some are also very important timber trees, producing wood of high quality. Several have edible seeds similar to pine nuts, and others produce valuable resin and amber. In the forests where they occur, they are usually dominant trees, often the largest species in the forest; the largest is Araucaria hunsteinii, reported to 89 m tall in New Guinea, with several other species reaching 50–65 m tall. A. heterophylla, the Norfolk Island pine, is a well-known landscaping and house plant from this taxon.
Skillful artisans in the Erzurum Province, Turkey, have used fossilized wood of Araucariaceae for centuries to manufacture jewelry and decorative items. It is known as "Oltustone", the name deriving from the town of Oltu, where it is most commonly excavated. Despite the fact that this semiprecious gemstone is classified as "stone", wood anatomy reveals it was fossilized pieces of trunks of Araucariacea. Oltustone, also called ‘Black Amber’ is unique to Turkey. It is dull and black, but when polished, acquires an attractive black sheen.<ref name="ReferenceA">Template:Cite journal</ref>
Fossil record
Fossils widely believed to belong to Araucariaceae include the form genera Araucarites (various), Agathoxylon and Araucarioxylon (wood), Brachyphyllum (leaves), Araucariacites and Dilwynites (pollen), and Protodammara (cones).
The oldest definitive records of Araucariaceae are from the Early Jurassic, though there are potential earlier Late Triassic records. Early representatives of Araucaria are widespread across both hemispheres by the Middle Jurassic, such as Araucaria mirabilis and Araucaria sphaerocarpa from the Middle Jurassic of Argentina and England respectively.<ref>Template:Cite journal</ref> The oldest records of the Wollemia-Agathis lineage from the Cretaceous, including Emwadea microcarpa from the Albian aged Winton Formation of Australia<ref>Template:Cite journal</ref> and Wairarapaia mildenhallii from the Albian-Cenomanian of New Zealand.<ref>Template:Cite journal</ref><ref name="Escapa 1153–1170"/> The oldest fossils currently confidently assignable to Agathis are those of Agathis immortalis from the Salamanca Formation of Patagonia, which dates to the Paleocene, approximately 64.67–63.49 million years ago. Agathis-like leaves are also known from the slightly older Lefipán Formation of the same region, which date to the very end of the Cretaceous.<ref>Template:Cite journal</ref> Araucariaceae fossils are also known from the latest Oligocene or earliest Miocene of the southwesternmost tip of Africa.<ref>Template:Cite journal</ref> Claimed records of Agathis from the Eocene of Canada based on chemical analysis of amber are questionable.<ref>Template:Cite journal</ref>
See also
References
Further reading
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