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Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the and periods, when it was distributed almost worldwide. Most of the Araucariaceae in the Northern Hemisphere vanished in the Cretaceous–Paleogene extinction event, and they are now largely confined to the Southern Hemisphere, except for a few species of in .


Description
Members of Araucariaceae are typically extremely tall trees, reaching heights of or more. They can also grow very large stem diameters; a tree ( Agathis australis) named Tāne Mahuta ("The Lord of the Forest") has been measured at tall with a diameter at breast height of . Its total wood volume is calculated to be , making it the third-largest conifer after Sequoia and (both from the Cupressaceae subfamily ).

The trunks are columnar and have relatively large with cortices. The branching is usually horizontal and tiered, arising regularly in whorls of three to seven branches or alternating in widely separated pairs.

(2023). 9780881929744, Timber Press. .

The leaves can be small, needle-like, and curved, or they can be large, broadly ovate, and flattened.

(1986). 9780521248594, Cambridge University Press. .
They are spirally arranged, persistent, and usually have parallel venation.

Like other conifers, they produce . Each tree can have both male and female cones () or they can have only male or female cones ().

Male cones are among the largest among all conifer cones, on average. They are cylindrical and drooping, somewhat resembling . They are borne singly on the tips of branches or the of leaves. They contain numerous arranged in whorls or spirals. Each has four to 20 elongated sacs attached to the lower surface at one end. The pollen grains are round and do not possess wings or air sacs.

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. The scales almost always bear only one seed on its upper surface, in contrast to two in true pines (family ). 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. 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 , about the size and weight of a large . They can drop from heights of .


Classification and genera
Araucariaceae is classified under the order , class of the division . The division includes all living . Recently however, some authorities treat Araucariaceae as a separate order, Araucariales. Araucariaceae contains three extant and about 41 species.
Jussieu

  • Araucaria angustifolia – Paraná pine
  • Araucaria araucana – monkey-puzzle or pehuén
  • Araucaria bidwilliibunya-bunya
  • Araucaria hunsteiniiklinki
  • Araucaria bernieri
  • Araucaria biramulata
  • Araucaria columnaris - Cook pine
  • Araucaria cunninghamii - Moreton Bay pine, hoop pine
  • Araucaria goroensis
  • Araucaria heterophylla – Norfolk Island pine
  • Araucaria humboldtensis
  • Araucaria laubenfelsii
  • Araucaria luxurians
  • Araucaria montana
  • Araucaria muelleri
  • Araucaria nemorosa
  • Araucaria schmidii
  • Araucaria scopulorum
  • Araucaria subulata
19 living species found in (where 13 species are endemic), , , , , , and .
Salisbury , Australia, , New Caledonia, Papua New Guinea, , , and the
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 of the Pinophyta based on of molecular data. It shows the position of Araucariaceae within the division.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

Relationships between living members of Araucariaceae.

Molecular evidence supports Araucariaceae and Podocarpaceae having diverged from each other during the late .


Distribution and habitat
Today, 41 species are known, in three genera: , and , 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 . In Malesia, Agathis extends a short distance into the Northern Hemisphere, reaching 18°N in the Philippines.


Uses
Several species are very popular ornamental trees in in subtropical regions, and some are also very important trees, producing of high quality. Several have edible seeds similar to , and others produce valuable and . In the 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 "", the name deriving from the town of , 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.


Fossil record
widely believed to belong to Araucariaceae include the form genera (various), and (wood), (leaves), and (pollen), and (cones).

The oldest definitive records of Araucariaceae are from the , though there are potential earlier Late Triassic records. Early representatives of Araucaria are widespread across both hemispheres by the , such as Araucaria mirabilis and Araucaria sphaerocarpa from the Middle Jurassic of Argentina and England respectively. The oldest records of the Wollemia- Agathis lineage from the , including from the aged of Australia and from the Albian- of New Zealand. The oldest fossils currently confidently assignable to Agathis are those of Agathis immortalis from the Salamanca Formation of Patagonia, which dates to the , 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.


See also


Further reading
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