{{Short description|Family of conifers}} {{Automatic taxobox | fossil_range = {{fossilrange|Early Jurassic|Present|earliest=Late Triassic}} (possible Late Triassic records) | image = Webysther 20190413132108 - Araucária (Araucaria angustifolia).jpg | image_caption = ''Araucaria angustifolia'' at Minas Gerais | taxon = Araucariaceae | authority = Henkel & W. Hochstetter | type_genus = ''Araucaria'' | type_genus_authority = Juss. | subdivision_ranks = Genera | subdivision = *''Agathis'' *''Araucaria'' *''Wollemia'' *†''Agathoxylon'' (wood, in part) *†''Araucarioides'' (leaves, ovulate cone) *†''Araucarites'' (ovulate cone) *†''Brachyphyllum'' (foliage, in part) *†''Emwadea'' (ovulate cone) *†''Pagiophyllum'' (foliage, in part) *†''Wairarapaia'' (ovulate cone) *†''Dilwynites'' (pollen) }}
'''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>{{Cite journal |last1=Stockey |first1=Ruth A. |last2=Rothwell |first2=Gar W. |date=July 2020 |title=Diversification of crown group Araucaria : the role of Araucaria famii sp. nov. in the Late Cretaceous (Campanian) radiation of Araucariaceae in the Northern Hemisphere |url=https://bsapubs.onlinelibrary.wiley.com/doi/10.1002/ajb2.1505 |journal=American Journal of Botany |language=en |volume=107 |issue=7 |pages=1072–1093 |doi=10.1002/ajb2.1505 |pmid=32705687 |issn=0002-9122}}</ref>
==Description== [[File:00 29 0496 Waipoua Forest NZ - Kauri Baum Tane Mahuta.jpg|thumb|upright=0.9|right|Tāne Mahuta ("Lord of the Forest"), a massive ''Agathis australis'' tree from New Zealand]] Members of Araucariaceae are typically extremely tall evergreen trees,<ref name="gym">{{cite web|url=http://www.conifers.org/ar/Araucariaceae.php|title=Araucariaceae|publisher=The Gymnosperm Database|access-date=November 19, 2011}}</ref> reaching heights of {{convert|60|m|ft|abbr=on}} or more.<ref name="ucmp">{{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}}</ref> They can also grow very large stem diameter. As an example of the family, the 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}},<ref name="kauri">{{cite web|url=http://www.conifers.org/ar/Agathis_australis.php|title=Agathis australis|publisher=The Gymnosperm Database|access-date=November 19, 2011}}</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">{{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 }}</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">{{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}}</ref>
The leaves can be small, needle-like, and curved, or they can be large, broadly ovate, and flattened.<ref name="walters">{{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}}</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">{{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}}</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">{{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}}</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. As an example of the size of cones in this plant family, the cones of the bunya bunya, ''Araucaria bidwillii'', weigh up to {{cvt|10|kg}},<ref name="TGD">{{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}}</ref> about the size and weight of a large pineapple. They can drop from heights of {{convert|23|m|ft|abbr=on}}.<ref name="ww"/>
==Classification and genera== [[File:AraucariaHfoliage.jpg|thumb|''Araucaria heterophylla'']] [[File:Agathisrobusta.JPG|thumb|''Agathis robusta'']] [[File:Wollemia nobilis fg03.JPG|thumb|''Wollemia nobilis'']] 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"/> thumb|1x1px Araucariaceae contains three extant genera and about 41 species.<ref name="wu"/> {| class="wikitable" |- ! Image !! Genus !! Living Species !! Distribution |- |175px |''Araucaria'' <small>Jussieu</small> |
* ''Araucaria angustifolia'' – Paraná pine * ''Araucaria araucana'' – monkey-puzzle or ''pehuén'' * ''Araucaria bernieri'' * ''Araucaria bidwillii'' – ''bunya-bunya'' * ''Araucaria biramulata'' * ''Araucaria columnaris'' – Cook pine * ''Araucaria cunninghamii'' – Moreton Bay pine, hoop pine * ''Araucaria goroensis'' * ''Araucaria heterophylla'' – Norfolk Island pine * ''Araucaria humboldtensis'' * ''Araucaria hunsteinii'' – ''klinki'' * ''Araucaria laubenfelsii'' * ''Araucaria luxurians'' * ''Araucaria montana'' * ''Araucaria muelleri'' * ''Araucaria nemorosa'' * ''Araucaria rulei'' * ''Araucaria schmidii'' * ''Araucaria scopulorum'' * ''Araucaria subulata'' |20 living species found in New Caledonia (where 14 species are endemic), Norfolk Island, Australia, New Guinea, Argentina, Chile, and Brazil. |- |175px |''Agathis'' <small>Salisbury</small> | * ''Agathis atropurpurea'' – black kauri, blue kauri (Queensland, Australia) * ''Agathis australis'' – kauri, New Zealand kauri (North Island, New Zealand) * ''Agathis borneensis'' (western Malesia, Borneo) * ''Agathis dammara'' (syn. ''A. alba'', ''A. celebica'', ''A. loranthifolia'') – Bindang (eastern Malesia) * ''Agathis flavescens'' (Peninsular Malaysia) * ''Agathis kinabaluensis'' (Borneo) * ''Agathis labillardierei'' (New Guinea) * ''Agathis lanceolata'' (New Caledonia) * ''Agathis lenticula'' (Borneo) * ''Agathis macrophylla'' (syn. ''A. vitiensis'') – Pacific kauri, dakua (Fiji, Vanuatu, Solomon Islands) * ''Agathis microstachya'' – bull kauri (Queensland, Australia) * ''Agathis montana'' (New Caledonia) * ''Agathis moorei'' – white kauri (New Caledonia) * ''Agathis orbicula'' (Borneo) * ''Agathis ovata'' (New Caledonia) * ''Agathis robusta'' – Queensland kauri (Queensland, Australia; New Guinea) * ''Agathis silbae'' (Vanuatu) | New Zealand, Australia, Vanuatu, New Caledonia, Papua New Guinea, Indonesia, Malaysia, and the Philippines |- |175px |''Wollemia'' <small>W.G. Jones, K.D. Hill & J.M. Allen</small> | *''Wollemia nobilis'' | 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> {{clade|style=line-height:100%; |1={{clade |1=Pinaceae |2={{clade |1={{clade |1='''Araucariaceae''' |2=Podocarpaceae }} |2={{clade |1=Sciadopityaceae |2={{clade |1=Cupressaceae |2={{clade |1=Cephalotaxaceae |2=Taxaceae }} }} }} }} }} }} Relationships between living members of Araucariaceae.<ref name="Escapa 1153–1170">{{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|bibcode=2013IJPlS.174.1153E |issn=1058-5893|hdl=11336/3583|s2cid=56238574|hdl-access=free}}</ref>
{{clade|{{clade |label1=Araucariaceae |1={{clade |1=''Araucaria'' |2={{clade |1=''Wollemia'' |2=''Agathis'' }} }} }}|style=font-size:100%;line-height:80%}} Molecular evidence supports Araucariaceae and Podocarpaceae having diverged from each other during the late Permian.<ref>{{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|url-access=subscription}}</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== {{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 {{Convert abbreviated|89|m}} tall in New Guinea, with several other species reaching {{Convert abbreviated|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">{{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 }}</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>{{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}}</ref> The oldest records of the ''Wollemia''-''Agathis'' lineage from the Cretaceous, including ''Emwadea microcarpa'' from the Albian aged Winton Formation of Australia<ref>{{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|url-access=subscription}}</ref> and ''Wairarapaia mildenhallii'' from the Albian-Cenomanian of New Zealand.<ref>{{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|bibcode=2006IJPlS.167.1259C |s2cid=85365035|issn=1058-5893|url-access=subscription}}</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>{{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 |bibcode=2018AmJB..105.1345E |hdl=11336/87592 |hdl-access=free }}</ref> Araucariaceae fossils are also known from the latest Oligocene or earliest Miocene of the southwesternmost tip of Africa.<ref>{{Cite journal |last1=Roberts |first1=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 |bibcode=2017GPC...150....1R |access-date=26 October 2024 |via=Elsevier Science Direct|hdl=2381/39417 |hdl-access=free }}</ref> Claimed records of ''Agathis'' from the Eocene of Canada based on chemical analysis of amber are questionable.<ref>{{Cite journal |last1=Archibald |first1=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).}}</ref>
==See also== *Paleobotany *Te Matua Ngahere ==References== {{Reflist|2}}
==Further reading== {{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 | issue = 1 | pages = 397–414 | doi = 10.1146/annurev.ecolsys.32.081501.114059 | bibcode = 2001AnRES..32..397K }} * {{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 }} {{Clear}}
{{Pinophyta}} {{Taxonbar|from=Q217697}} {{Authority control}}
Category:Araucariaceae Category:Pinales families Category:Extant Hettangian first appearances Category:Araucariales