Pinales
Pinales | |
---|---|
Pinus sylvestris | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Gymnospermae |
Division: | Pinophyta |
Class: | Pinopsida |
Subclass: | Pinidae |
Order: | Pinales Gorozh.[1][2] |
Type genus | |
Pinus
| |
Families | |
(approximate number of species in parentheses) | |
Synonyms | |
The
Multiple molecular studies indicate this order being paraphyletic with respect to Gnetales, with studies recovering Gnetales as either a sister group to Pinaceae or being more derived than Pinaceae but sister to the rest of the group.
Taxonomy
History
- Cycadeaceae (cycads)
- Pinaceae (conifers)
- Taxaceae (taxads)
- Gnetaceae
In contrast,
- Gnetaceae
- Coniferae
- Cycadaceae
In the Engler system (1903) Gymnospermae is listed as a subdivision (Unterabteilung) and adopted more of a splitter approach, including extinct taxa, with the following six classes;[12]
- Cycadales
- Bennettitales
- Cordaitales
- Ginkgoales
- Coniferae
- Gnetales
During this period,
- Cycadoideae
- Peucideae(Coniferae)
A system of two groups was maintained by the most commonly used classification in the twentieth century,[13] the revision of the Engler system by Pilger (1926), who grouped 12 families of the Gymnospermae subdivision into 2 classes;[14]
- Coniferales (Coniferae)
- Gnetales
The treatment of Gymnosperms as two groups, though with varying composition and names, was followed for most of the twentieth century, including the systems of Chamberlain (1935),[15] Benson (1957)[16] and Cronquist (1960).[17]
In the latter, Cronquist divided Gymnospermae into two divisions;
- division Coniferophyta
- class Coniferae
- class Chlamydospermae(Gnetales)
- division Cycadophyta
- class Cycadae
Benson,(1957)[16] who introduced the term Pinales, divided gymnosperms into four classes;[9]
- Conopsida (conifers, including Pinales)
- Ephedropsida
- Gnetopsida
- Cycadopsida
In a later revision, in collaboration with two other taxonomists (1966), Cronquist merged all the gymnosperms into a single division, Pinophyta, with three subdivisions reflecting the main lineages;[18][9]
- Cycadicae
- Pinicae
- Gneticae
In the era of molecular phylogenetics, De-Zhi and colleagues (2004) once again proposed a division of 12 gymnosperm families into two classes;[9]
- Cycadopsida
- order Cycadales
- Coniferopsida
- subclass Multinervidae (6 orders)
- subclass Taxidae
- order Taxales
- order Pinales
With the development of the
- subclass Cycadidae Pax(1)
- subclass Ginkgooidae Engl. (1)
- subclass Gnetidae Pax (3)
- subclass Pinidae Zimmerm.(conifers) (1)
Controversies
Phylogeny
Gymnosperms form a group of four subclasses among the
There are about 1000 extant gymnosperm
- Cycadidae (Cycadophyta, cycads 10, 300)
- Ginkgoidae (Ginkgophyta, ginkgo 1, 1)
- Gnetidae (Gnetophyta 3, 100)
- Pinidae (Pinophyta,[18][28] conifers 70, 600)
The term Pinophyta has also been used to include all conifers, extinct and extant, with Pinales representing all the extant conifers.[29]
- Pinales Gorozh. (Pinaceae)
- Araucariales Gorozh. (Araucariaceae, Podocarpaceae)
- )
However, the exact phylogeny remained a topic that was 'hotly debated", in particular whether the main lineages were best represented by the four subclasses of Christenhusz and colleagues or the more traditional five clades (cycads, ginkgos, cupressophytes, Pinaceae and gnetophytes).[27] In 2014 the first complete molecular phylogeny was published, based on 90 species representing all extant genera. This established cycads as the basal group, followed by Ginkgoaceae, as sister to the remaining gymnosperms, and supporting the ‘gnepine’ hypothesis. This analysis favours the five clade hypothesis, the remaining clade following divergence of the Pinidae, are referred to as the conifer II clade, or cupressophytes, in distinction from the conifer I clade (Gnetidae, Pinidae).[30] This clade, in turn, has two lineages. The first consisting of Sciadopityaceae and the Araucariales, the second being the Cupressales. In the Christenhusz scheme, the Sciadopityaceae were considered to be within Cupressales. The term Cupressaceae s.l. refers to the inclusion of Taxodiaceae.[31] These relationships are shown in this cladogram, although no formal taxonomic revision was undertaken.[31]
Phylogeny of Gymnosperms, subclasses, orders, families[31][2] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
A more comprehensive analysis was undertaken by Ran and colleagues in 2018, as part of a detailed phylogeny of all seed plants.[32] This forms the basis of the Tracheophyte Phylogeny Poster[30] and the Angiosperm Phylogeny Website.[33]
Subdivision
Historically conifers, in the order Pinales have been considered to consist of six to seven extant families, based on the classification of class Coniferae by
- Araucariaceae
- redwood)
- )
- Podocarpaceae
- yews)
- Cephalotaxaceae
- Taxodiaceae
Subsequent revisions merged the Taxodiaceae and Cupressaceae, and placed
Christenhusz and colleagues (2011) included only one family in Pinales, Pinaceae,[2] a practice subsequently followed by the Angiosperm Phylogeny Website[33] and the Gymnosperm Database.[35] In this restricted model Pinales (Pinaceae) comprisea 11 genera and about 225 species, all of the other conifers originally included in this order, being included in other orders such as Cupressales.[2]
Notes
- ^ Taxon names beginning with the root conifer- are considered illegitimate because they are not based on an underlying genus[2]
- Linnean Societyin 1825, published in 1826
- ^ Phanerogam, or seed plant, indicating visible sexual organs, and gymnosperm indicating exposed seeds
- ^ Angiosperm indicating enclosed seeds
- Equisetopsidasensu stricto when used as a class of ferns, synonymous with Equisetidae
References
- ^ a b Gorozhankin 1904.
- ^ a b c d e f g h Christenhusz et al 2011.
- ^ a b c Eckenwalder 2008.
- ^ a b Gernandt et al 2011.
- ^ Brown 1826.
- ^ Tomlinson 2012.
- ^ Brongniart 1828–1837.
- ^ Lindley 1830.
- ^ a b c d De-Zhi et al 2004.
- ^ Lindley 1853.
- ^ Bentham & Hooker 1862–1883.
- ^ Engler 1903.
- ^ a b Brunsfield et al 1994.
- ^ a b Pilger 1926.
- ^ Chamberlain 1935.
- ^ a b Benson 1957.
- ^ Cronquist 1960.
- ^ a b Cronquist et al 1966.
- ^ a b c Chase & Reveal 2009.
- ^ Farjon 2001.
- ^ a b Nagalingum & Burleigh 2018.
- ^ Eckenwalder 2009.
- ^ Farjon 2010.
- ^ Debreczy & Rácz 2011.
- ^ Farjon & Filer 2013.
- ^ Farjon 2018.
- ^ a b c d Wang & Ran 2014.
- ^ Reveal 1995.
- ^ Delevoryas 2015.
- ^ a b Cole & Bachelier 2019.
- ^ a b c Lu et al 2014.
- ^ Ran et al 2018.
- ^ a b Stevens 2020.
- ^ Davy 1937.
- ^ a b Earle 2020.
Bibliography
Books
- Anderson, John Malcolm; Anderson, Heidi M.; Cleal, Christopher J. (2007). Brief History of the Gymnosperms: Classification, Biodiversity, Phytogeography and Ecology. ISBN 978-1-919976-39-6.
- Benson, Lyman David (1957). Plant Classification. Heath.
- Benson, Lyman David (1979). Plant Classification (2nd) ed.). Heath. ISBN 978-0-669-01489-1.
- Benson, Lyman David (1979). Plant Classification (2nd) ed.). Heath.
- Bentham, G.; Hooker, J.D. (1862–1883). "Gymnospermeae". Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita (3 vols.) (in Latin). Vol. 3(1) 1880. London: L Reeve & Co. pp. 417–447.
- Brongniart, Adolphe (1828–1837). Histoire des végétaux fossiles, ou, Recherches botaniques et géologiques sur les végétaux renfermés dans les diverses couches du globe 2 vols. Vol. i 1828. Paris: G. Dufour et Ed. d'Ocagne. p. 22.
- Eckenwalder, James E. (2009). Conifers of the World: The Complete Reference. ISBN 978-0-88192-974-4.
- Gebrüder BorntraegerVerlag. pp. 70–75.
- Prantl, Karl, eds. (1924) [1887–1915]. Die natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten (in German). Vol. 21 parts (Abteilungen) (2nd ed.). Berlin: Duncker & Humblot., see also Die Natürlichen Pflanzenfamilien
- Pilger, Robert (1926). Klasse Coniferae. pp. 121–403., in Engler & Prantl (1924)vol. 13 Gymnospermae
- Debreczy, Zsolt; Rácz, István (2011). Conifers Around the World: Conifers of the Temperate Zones and Adjacent Regions. DendroPress. ISBN 978-963-219-061-7.
- ISBN 978-1-84246-025-2.
- — (2010). A Handbook of the World's Conifers (2 Vols.). ISBN 978-90-04-17718-5.
- —; Filer, Denis (2013). An Atlas of the World's Conifers: An Analysis of their Distribution, Biogeography, Diversity and Conservation Status. ISBN 978-90-04-21181-0.
- Gorozhankin, Ivan Nikolaevich [in Russian] (1904). Лекции по морфологии и систематики архегониальных растений (Lekts. Morf. Sist. Archegon. Lectures on the morphology and taxonomy of archegonial plants) (in Russian). Moscow: Русская" типо-литография. p. 88., see also Горожанкин, Иван Николаевич (1895). Морфология и систематика архегониальных растений: лекции профессора И.Н. Горожанкина 1894-95 г. Gymnaspermae (Archispermae). Часть III. "Русская" типо-литография.
- Lindley, John (1830). "General view". An introduction to the natural system of botany: or, A systematic view of the organisation, natural affinities, and geographical distribution, of the whole vegetable kingdom: together with the uses of the most important species in medicine, the arts, and rural or domestic economy (1st ed.). London: Longman. p. xxxv.
- Lindley, John (1853) [1846]. "Gymnogens". The Vegetable Kingdom: or, The structure, classification, and uses of plants, illustrated upon the natural system (3rd. ed.). London: Bradbury & Evans. pp. 221–234.
- Plomion, Christophe; Bousquet, Jean; Kole, Chittaranjan, eds. (2011). Genetics, Genomics and Breeding of Conifers. ISBN 978-1-57808-719-8.
- Gernandt, David S; Willyard, Ann; Syring, John V; Liston, Aaron (2011). The Conifers (Pinophyta). pp. 1–39., in Plomion et al (2011)
Encyclopaedias
- Delevoryas, T (13 April 2015). "Gymnosperm". Encyclopædia Britannica. Retrieved 6 August 2020.
- Eckenwalder, James Emory (3 September 2008). "Conifer". Encyclopædia Britannica. Retrieved 27 July 2020.
Articles
- .
- Brunsfeld, Steven J.; Soltis, Pamela S.; Soltis, Douglas E.; Gadek, Paul A.; Quinn, Christopher J.; Strenge, Darren D.; Ranker, Tom A. (April 1994). "Phylogenetic Relationships Among the Genera of Taxodiaceae and Cupressaceae: Evidence from rbcL Sequences". JSTOR 2419600.
- Chamberlain, Charles Joseph (June 1935). "The Gymnosperms" (PDF). S2CID 39793752.
- .
- S2CID 86797396.
- S2CID 43144314.
- JSTOR 1217531.
- Davy, J. Burtt (1937). "The classification of Coniferae II". Forestry: An International Journal of Forest Research. 11 (2): 122–123. .
- De-Zhi, Fu; Yong, Yang; Guang-Hua, Zhu (2004). "A New Scheme of Classification of Living Gymnosperms at Family Level". JSTOR 4111081.
- De La Torre, Amanda R.; Piot, Anthony; Liu, Bobin; Wilhite, Benjamin; Weiss, Matthew; Porth, Ilga (January 2020). "Functional and morphological evolution in gymnosperms: A portrait of implicated gene families". Evolutionary Applications. 13 (1): 210–227. PMID 31892953.
- .
- Lu, Ying; Ran, Jin-Hua; Guo, Dong-Mei; Yang, Zu-Yu; Wang, Xiao-Quan (15 September 2014). "Phylogeny and Divergence Times of Gymnosperms Inferred from Single-Copy Nuclear Genes". PMID 25222863.
- Ran, Jin-Hua; Shen, Ting-Ting; Wang, Ming-Ming; Wang, Xiao-Quan (27 June 2018). "Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms". PMID 29925623.
- Ran, Jin-Hua; Shen, Ting-Ting; Wu, Hui; Gong, Xun; Wang, Xiao-Quan (December 2018). "Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis". S2CID 52110440.
- Reveal, James L (August 1995). "Newly required suprageneric names in Magnoliophyta". Phytologia. 79 (2): 68–76.
- Tomlinson, P. B. (2012). "Rescuing Robert Brown—The Origins of Angio-Ovuly in Seed Cones of Conifers". S2CID 5403322.
- Wang, Xiao-Quan; Ran, Jin-Hua (June 2014). "Evolution and biogeography of gymnosperms". S2CID 4662571.
Websites
- Cole, Theodor C. H.; Bachelier, Julien B (October 2019). "Tracheophyte Phylogeny Poster - Vascular Plants: Systematics and Characteristics, 2019". Dahlem Centre of Plant Sciences (DCPS), Institute of Biology – Botany, Freie Universität Berlin. Retrieved 24 August 2020.
- Stevens, P.F. (2020) [2001]. "Pinales". AP Web v. 14. Missouri Botanical Garden. Retrieved 28 July 2020. (see also Angiosperm Phylogeny Website)
- Earle, Christopher J. (2020). "The Gymnosperm Database".
- Nagalingum, Nathalie; Burleigh, Gordon (2018). "The Gymnosperm Phylogeny Group". Botany 2018 (Abstract). Botanical Society of America. Retrieved 30 July 2020.
- Eckenwalder, James E (2007). "Gymnosperm classification". Flora of North America. Volume 1, Chapter 13. Retrieved 11 August 2020.