Araucariaceae

File:00 29 0496 Waipoua Forest NZ - Kauri Baum Tane Mahuta.jpg ("Lord of the Forest"), a massive Agathis australis tree from New Zealand]]

Members of Araucariaceae are typically extremely tall evergreen trees,{{cite web|url=http://www.conifers.org/ar/Araucariaceae.php|title=Araucariaceae|publisher=The Gymnosperm Database|access-date=November 19, 2011}} reaching heights of {{convert|60|m|ft|abbr=on}} or more.{{cite web|url=http://www.conifers.org/ar/Araucariaceae.php|title=Araucariaceae: life history and ecology|publisher=University of California Museum of Paleontology|access-date=November 19, 2011}} 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 {{convert|45.2|m|ft|abbr=on}} tall with a diameter at breast height of {{convert|491|cm|ft|abbr=on}}. Its total wood volume is calculated to be {{convert|516.7|m3|cuft|abbr=on}},{{cite web|url=http://www.conifers.org/ar/Agathis_australis.php|title=Agathis australis|publisher=The Gymnosperm Database|access-date=November 19, 2011}} making it the third-largest conifer after Sequoia and Sequoiadendron (both from the Cupressaceae subfamily Sequoioideae).

The trunks are columnar and have relatively large piths with resinous cortices.{{cite journal |author1=Fu Liguo |author2=Li Nan |author3=Robert R. Mill |title=Araucariaceae |journal=Flora of China |volume=4 |pages=9–10 |year=1999 |url=http://hua.huh.harvard.edu/china/mss/volume04/ARAUCARIACEAE.published.pdf |access-date=2011-11-19 |url-status=dead |archive-url=https://web.archive.org/web/20110524173027/http://hua.huh.harvard.edu/china/mss/volume04/ARAUCARIACEAE.published.pdf |archive-date=May 24, 2011 }} The branching is usually horizontal and tiered, arising regularly in whorls of three to seven branches or alternating in widely separated pairs.{{cite book|author=James E. Eckenwalder|title =Conifers of the world: the complete reference|publisher =Timber Press|year =2009|page=70|isbn =978-0-88192-974-4|url =https://books.google.com/books?id=b9aqcxSqYCkC&pg=PA70}}

The leaves can be small, needle-like, and curved, or they can be large, broadly ovate, and flattened.{{cite book|author=Stuart Max Walters|title =The European Garden Flora: Pteridophyta, Gymnospermae, Angiospermae|publisher =Cambridge University Press|year =1986|page=72|isbn =978-0-521-24859-4|url =https://books.google.com/books?id=1dd5M-ToXAcC&pg=PA72}} They are spirally arranged, persistent, and usually have parallel venation.

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).{{cite web

|url=http://www.botany.hawaii.edu/faculty/carr/araucari.htm|title=Araucariaceae|author=Gerald Carr|publisher=University of Hawaii|access-date=November 19, 2011}}

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.

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 Pinaceae).{{cite web|url=http://waynesword.palomar.edu/ecoph27.htm|title=The Araucaria Family: Araucariaceae|author=Wayne P. Armstrong|publisher=Wayne's Word, Paloma College|access-date=November 19, 2011|archive-date=December 3, 2011|archive-url=https://web.archive.org/web/20111203031105/http://waynesword.palomar.edu/ecoph27.htm|url-status=dead}} 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 {{cvt|10|kg}},{{cite web |title=Araucaria biwillii (Bunya pine) description |url=https://www.conifers.org/ar/Araucaria_bidwillii.php |website=The Gymnosperm Database |access-date=20 January 2023}} about the size and weight of a large pineapple. They can drop from heights of {{convert|23|m|ft|abbr=on}}.

File:AraucariaHfoliage.jpg]]

File:Agathisrobusta.JPG]]

File:Wollemia nobilis fg03.JPG]]

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.

File:WollemiaNobilisPineKiefer.jpg

Araucariaceae contains three extant genera and about 41 species.

Below is the phylogeny of the Pinophyta based on cladistic analysis 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

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Relationships between living members of Araucariaceae.{{Cite journal|last1=Escapa|first1=Ignacio H.|last2=Catalano|first2=Santiago A.|date=October 2013|title=Phylogenetic Analysis of Araucariaceae: Integrating Molecules, Morphology, and Fossils|url=https://www.journals.uchicago.edu/doi/10.1086/672369|journal=International Journal of Plant Sciences|language=en|volume=174|issue=8|pages=1153–1170|doi=10.1086/672369|issn=1058-5893|hdl=11336/3583|s2cid=56238574|hdl-access=free}}

{{clade|{{clade

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|1={{clade

|1=Araucaria

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|1=Wollemia

|2=Agathis

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Molecular evidence supports Araucariaceae and Podocarpaceae having diverged from each other during the late Permian.{{Cite journal|last1=Stull|first1=Gregory W.|last2=Qu|first2=Xiao-Jian|last3=Parins-Fukuchi|first3=Caroline|last4=Yang|first4=Ying-Ying|last5=Yang|first5=Jun-Bo|last6=Yang|first6=Zhi-Yun|last7=Hu|first7=Yi|last8=Ma|first8=Hong|last9=Soltis|first9=Pamela S.|last10=Soltis|first10=Douglas E.|last11=Li|first11=De-Zhu|date=July 19, 2021|title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms|url=https://www.nature.com/articles/s41477-021-00964-4|journal=Nature Plants|language=en|volume=7|issue=8|pages=1015–1025|doi=10.1038/s41477-021-00964-4|pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |issn=2055-0278}}

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.

{{Further|Agathis#Uses|Araucaria#Uses|Wollemia#Uses}}

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.{{cite journal | last1 = Kutluk | display-authors = etal | year = 2012 | title = First Report of Araucariaceae wood (Agathoxylon sp.) from the Late Cretaceous of Turkey| journal = IAWA Journal | volume = 33 | issue = 3| pages = 319–326 | doi=10.1163/22941932-90000097| doi-access = free }}

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.{{Cite journal|last1=Leslie|first1=Andrew B.|last2=Beaulieu|first2=Jeremy|last3=Holman|first3=Garth|last4=Campbell|first4=Christopher S.|last5=Mei|first5=Wenbin|last6=Raubeson|first6=Linda R.|last7=Mathews|first7=Sarah|date=September 2018|title=An overview of extant conifer evolution from the perspective of the fossil record|journal=American Journal of Botany|language=en|volume=105|issue=9|pages=1531–1544|doi=10.1002/ajb2.1143|pmid=30157290|doi-access=free}} The oldest records of the Wollemia-Agathis lineage from the Cretaceous, including Emwadea microcarpa from the Albian aged Winton Formation of Australia{{Cite journal|last1=Dettmann|first1=Mary E.|last2=Clifford|first2=H. Trevor|last3=Peters|first3=Mark|date=June 2012|title=Emwadea microcarpa gen. et sp. nov.—anatomically preserved araucarian seed cones from the Winton Formation (late Albian), western Queensland, Australia|url=http://www.tandfonline.com/doi/abs/10.1080/03115518.2012.622155|journal=Alcheringa: An Australasian Journal of Palaeontology|language=en|volume=36|issue=2|pages=217–237|doi=10.1080/03115518.2012.622155|bibcode=2012Alch...36..217D |s2cid=129171237|issn=0311-5518}} and Wairarapaia mildenhallii from the Albian-Cenomanian of New Zealand.{{Cite journal|last1=Cantrill|first1=David J.|last2=Raine|first2=J. Ian|date=November 2006|title=Wairarapaia mildenhallii gen. et sp. nov., a New Araucarian Cone Related to Wollemia from the Cretaceous (Albian-Cenomanian) of New Zealand|url=http://www.journals.uchicago.edu/doi/10.1086/507608|journal=International Journal of Plant Sciences|language=en|volume=167|issue=6|pages=1259–1269|doi=10.1086/507608|s2cid=85365035|issn=1058-5893}} 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.{{Cite journal |last1=Escapa |first1=Ignacio H. |last2=Iglesias |first2=Ari |last3=Wilf |first3=Peter |last4=Catalano |first4=Santiago A. |last5=Caraballo-Ortiz |first5=Marcos A. |last6=Rubén Cúneo |first6=N. |date=August 2018 |title=Agathis trees of Patagonia's Cretaceous-Paleogene death landscapes and their evolutionary significance |journal=American Journal of Botany |language=en |volume=105 |issue=8 |pages=1345–1368 |doi=10.1002/ajb2.1127 |pmid=30074620 |s2cid=51908977 |issn=0002-9122|doi-access=free |hdl=11336/87592 |hdl-access=free }} Araucariaceae fossils are also known from the latest Oligocene or earliest Miocene of the southwesternmost tip of Africa.{{Cite journal |last=Roberts |first=D.L. |last2=Neumann |first2=F.H. |last3=Cawthra |first3=H.C. |last4=Carr |first4=A.S. |last5=Scott |first5=L. |last6=Durugbo |first6=E.U. |last7=Humphries |first7=M.S. |last8=Cowling |first8=R.M. |last9=Bamford |first9=M.K. |last10=Musekiwa |first10=C. |last11=MacHutchon |first11=M. |date=March 2017 |title=Palaeoenvironments during a terminal Oligocene or early Miocene transgression in a fluvial system at the southwestern tip of Africa |url=https://www.sciencedirect.com/science/article/abs/pii/S0921818116303022 |journal=Global and Planetary Change |language=en |volume=150 |pages=1–23 |doi=10.1016/j.gloplacha.2017.01.007 |access-date=26 October 2024 |via=Elsevier Science Direct|hdl=2381/39417 |hdl-access=free }} Claimed records of Agathis from the Eocene of Canada based on chemical analysis of amber are questionable.{{Cite journal |last=Archibald |first=S. Bruce |last2=Makarkin |first2=Vladimir N. |date=February 2004 |title=New genus of minute Berothidae (Neuroptera) from Early Eocene amber of British Columbia |url=https://citeseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=cc1655f5fd38e085842b2d4fe15b0378892e050a |journal=The Canadian Entomologist |language=en |volume=136 |issue=1 |pages=61–76 |doi=10.4039/n03-043 |issn=0008-347X |quote="Nuclear magnetic resonance (NMR) spectroscopy indicated that the amber was formed by resin from the genus Agathis (Araucariaceae) (Poinar et al. 1999); however, NMR spectroscopy indicating the origins of various Cretaceous ambers from Araucariaceae may be problematic (Grimaldi et al. 2000).}}

• Paleobotany
• Te Matua Ngahere

{{Reflist|2}}

{{Commons category|Araucariaceae}}

{{Wikispecies|Araucariaceae}}

• {{cite journal | last1 = Cookson | first1 = I. C. | last2 = Duigan | first2 = S. L. | year = 1951 | title = Tertiary Araucariaceae from South-eastern Australia, with notes on living species | url =http://www.publish.csiro.au/bi/pdf/bi9510415 | journal = Australian Journal of Scientific Research Series B (Biological Sciences) | volume = 4 | pages = 415–449 }}
• {{cite journal | last1 = Kendall | first1 = Mabel W | year = 1949 | title = A Jurassic member of the Araucariaceae | journal = Annals of Botany |series=New Series | volume = 13 | issue = 50| pages = 151–161 | doi = 10.1093/oxfordjournals.aob.a083211 }}
• {{cite journal | last1 = Kershaw | first1 = Peter | last2 = Wagstaff | first2 = Barbara | year = 2001 | title = The Southern Conifer Family Araucariaceae: History, Status, and Value for Paleoenvironmental Reconstruction | journal = Annual Review of Ecology and Systematics | volume = 32 | pages = 397–414 | doi = 10.1146/annurev.ecolsys.32.081501.114059 }}
• {{cite journal | last1 = Krasilov | first1 = Valentin A | year = 1978 | title = Araucariaceae as indicators of climate and paleolatitudes | doi = 10.1016/0034-6667(78)90008-8 | journal = Review of Palaeobotany and Palynology | volume = 26 | issue = 1–4| pages = 113–124 | bibcode = 1978RPaPa..26..113K }}
• {{cite journal | last1 = Pye | first1 = Matthew G. | last2 = Henwood | first2 = Murray J. | last3 = Gadek | first3 = Paul A. | year = 2009 | title = Differential levels of genetic diversity and divergence among populations of an ancient Australian rainforest conifer, Araucaria cunninghamii | journal = Plant Systematics and Evolution | volume = 277 | issue = 3/4| pages = 173–185 | doi = 10.1007/s00606-008-0120-1 | bibcode = 2009PSyEv.277..173P | s2cid = 21846658 }}
• {{cite journal | last1 = Setoguchi | first1 = Hiroaki | display-authors = etal | year = 1998 | title = Phylogenetic relationships within Araucariaceae based on rbcL gene sequences | journal = American Journal of Botany | volume = 85 | issue = 11 | pages = 1507–1516 | doi = 10.2307/2446478 | pmid = 21680310 | jstor = 2446478 | doi-access = free }}
• {{cite journal | last1 = Stockey | first1 = Ruth A | year = 1982 | title = The Araucariaceae: an evolutionary perspective | journal = Review of Palaeobotany and Palynology | volume = 37 | issue = 1–2| pages = 133–154 | doi=10.1016/0034-6667(82)90041-0| bibcode = 1982RPaPa..37..133S }}
• {{cite journal | last1 = Stockey | first1 = Ruth A | year = 1994 | title = Mesozoic Araucariaceae: morphology and systematic relationships | journal = Journal of Plant Research | volume = 107 | issue = 4| pages = 493–502 | doi = 10.1007/BF02344070 | bibcode = 1994JPlR..107..493S | s2cid = 20148157 }}

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{{Pinophyta}}

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Category:Pinales families

Category:Extant Hettangian first appearances

Category:Araucariales