Pinales

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Pinales
Scots pine, Crow Wood. - geograph.org.uk - 139303.jpg
Pinus sylvestris
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Pinophyta
Class: Pinopsida
Subclass: Pinidae
Order: Pinales
Gorozh. [1] [2]
Type genus
Pinus
L.
Families

(approximate number of species in parentheses)

Synonyms

The order Pinales in the division Pinophyta, class Pinopsida, comprises all the extant conifers. The distinguishing characteristic is the reproductive structure known as a cone produced by all Pinales. All of the extant conifers, such as Araucaria, cedar, celery-pine, cypress, fir, juniper, kauri, larch, pine, redwood, spruce, and yew, are included here. Some fossil conifers, however, belong to other distinct orders within the division Pinophyta.

Contents

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

Brown (1825) [lower-alpha 2] [5] first discerned that there were two groups of seed plants, distinguished by the form of seed development, based on whether the ovules were exposed, receiving pollen directly, or enclosed, which do not. [6] Shortly afterwards, Brongniart (1828) coined the term Phanérogames gymnosperms [lower-alpha 3] to describe the former group. [7] The distinction was then formalized by Lindley (1830), dividing what he referred to as the subclass Dicotyledons into two tribes, Gymnosperms and Angiosperms. [lower-alpha 4] In the gymnosperms (or Gymnospermae) Lindley included two orders, the Cycadeae and the Coniferae. [8] [9] In his final work (1853) he described Gymnogens as a class with four orders; [10]

In contrast, Bentham and Hooker (1880) included only three orders in the class Gymnospermeae, by including taxads within Coniferae; [11]

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]

During this period, Gorozhankin published his treatise on Gymnosperms (1895), for which he bears the botanical authority for Pinales, Gorozh.. In his classification, Gymnospermae (alternatively named Archespermae) was a class of the division Archegoniatae, divided into subclasses; [1]

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]

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;

Benson,(1957) [16] who introduced the term Pinales, divided gymnosperms into four classes; [9]

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]

In the era of molecular phylogenetics, De-Zhi and colleagues (2004) once again proposed a division of 12 gymnosperm families into two classes; [9]

With the development of the Angiosperm Phylogeny Group came a major realignment of the linear classification of the land plants, by Chase and Reveal (2009). In this system, the land plants form a class, Equisetopsida s.l. (sensu lato) or sensu Chase & Reveal, [lower-alpha 5] also known as embryophytes or Embryophyceae nom. illeg. . [19] Class Equisetopsida s.l. is divided into 14 subclades as subclasses, including Magnoliidae (angiosperms). The gymnosperms are represented by four of these subclasses, placing them in a sister group relationship to angiosperms. Subclasses (number of orders); [19]

Controversies

Gymnosperm (Acrogymnospermae) taxonomy has been considered controversial, and lacks consensus. [20] [3] [21] As taxonomic classification transformed from being based solely on plant morphology to molecular phylogenetics, the number of taxonomic publications increased considerably after 2008, [22] [23] [2] [24] [25] however, these approaches have not been uniform. A taxonomic classification has been complicated by the relationship of extant to extinct taxa, and within extinct taxa, and particularly the placement of Gnetophyta. The latter have been variously classified as basal to all gymnosperms, sister group to conifers (‘gnetifer’ hypothesis) or sister to Pinaceae (‘gnepine’ hypothesis) in which they are classified within the conifers. [26] The extant conifers most likely form a monophyletic group. [4] [2] In 2018, the Gymnosperm Phylogeny Group was established, analogous to the Angiosperm Phylogeny Group and Pteridophyte Phylogeny Group, with the intention of reaching a consensus. [21]

Phylogeny

Gymnosperms form a group of four subclasses among the spermatophytes (seed bearing plants). In turn, the seed plants together with the monilophyte fern subclasses make up the tracheophytes (vascular plants), part of the class Equisetopsida (embryophytes or land plants), as opposed to the green algae. Among the seed plants, the gymnosperms are a sister group to the subclass Magnoliidae (angiosperms or flowering plants).

There are about 1000 extant gymnosperm species, distributed over about 12 families and 83 genera. Many of these genera are monotypic (41%), and another 27% are oligotypic (2–5 species). [27] The four subclasses have also been treated as divisions of the Spermatophytes. Alternative names and the approximate number of genera and species in each are; [27]

The term Pinophyta has also been used to include all conifers, extinct and extant, with Pinales representing all the extant conifers. [29]

Christenhusz and colleagues extended the system of Chase and Reveal [19] to provide a revised classification of gymnosperms in 2011, based on the above four subclades. [2] In this scheme, the Pinidae comprise three orders, including Pinales, and 6 families;

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]
Gymnosperms

Cycadidae (Cycadales)

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 Pilger (1926), considered the standard through most of the twentieth century. [13] These families were treated as a single order, in distinction to some earlier systems. [34] His families were; [14]

Subsequent revisions merged the Taxodiaceae and Cupressaceae, and placed Sciadopitys , formerly in Cupressaceae, into a separate family (Sciadopityaceae). [35] Cephalotaxaceae had previously been recognized as a separate family, but was subsequently included in Taxaceae. Similarly Phyllocladaceae were included in Podocarpaceae. Yews (Taxaceae) have sometimes been treated as a separate order (Taxales). [27]

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

  1. Taxon names beginning with the root conifer- are considered illegitimate because they are not based on an underlying genus [2]
  2. Read before the Linnean Society in 1825, published in 1826
  3. Phanerogam, or seed plant, indicating visible sexual organs, and gymnosperm indicating exposed seeds
  4. Angiosperm indicating enclosed seeds
  5. This term should not be confused with Equisetopsida sensu stricto when used as a class of ferns, synonymous with Equisetidae

Related Research Articles

<span class="mw-page-title-main">Conifer</span> Group of cone-bearing seed plants

Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta, also known as Coniferophyta or Coniferae. The division contains a single extant class, Pinopsida. All extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 2002, Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species.

<span class="mw-page-title-main">Araucariaceae</span> Family of plants

Araucariaceae – also known as araucarians – is a family of coniferous trees, with three living genera, Araucaria, Agathis, and Wollemia. While the family was distributed globally during the Jurassic and Cretaceous periods, in their native distribution they are now largely confined to the Southern Hemisphere, except for a few species of Agathis in Southeast Asia.

<span class="mw-page-title-main">Gnetophyta</span> Division of plants containing three genera of gymnosperms

Gnetophyta is a division of plants, grouped within the gymnosperms, that consists of some 70 species across the three relict genera: Gnetum, Welwitschia, and Ephedra. The earliest unambiguous records of the group date to the Jurassic, and they achieved their highest diversity during the Early Cretaceous. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of small tubes (xylem) that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question, with many recent phylogenies finding them to be nested within the conifers.

<span class="mw-page-title-main">Taxales</span> Order of conifers

The plant order Taxales was until recently treated as a distinct order in the division Pinophyta, class Pinopsida, and included only those species in the family Taxaceae, known commonly as yews. Under this interpretation, all other conifers were classified separately in the order Pinales. Recent genetic and micromorphological studies, however, have shown the Taxaceae are closely related to the other conifers, particularly so to the family Cephalotaxaceae. The order Taxales is therefore no longer recognised as distinct, and the family Taxaceae is now included in the order Cupressales. See Pinophyta for more details.

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms are a group of seed-producing plants that includes conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, Ginkgo. Gymnosperm lifecycles involve alternation of generations. They have a dominant diploid sporophyte phase and a reduced haploid gametophyte phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

<span class="mw-page-title-main">Pinaceae</span> Family of conifers

The Pinaceae, or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, firs, hemlocks, piñons, larches, pines and spruces. The family is included in the order Pinales, formerly known as Coniferales. Pinaceae have distinctive cones with woody scales bearing typically two ovules, and are supported as monophyletic by both morphological trait and genetic analysis. They are the largest extant conifer family in species diversity, with between 220 and 250 species in 11 genera, and the second-largest in geographical range, found in most of the Northern Hemisphere, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of boreal, coastal, and montane forests. One species, Pinus merkusii, grows just south of the equator in Southeast Asia. Major centres of diversity are found in the mountains of southwest China, Mexico, central Japan, and California.

<span class="mw-page-title-main">Pteridophyte</span> Group of plants that reproduce by spores

A pteridophyte is a vascular plant that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden.

<i>Torreya</i> Genus of conifers

Torreya is a genus of conifers comprising six or seven species placed in the family Taxaceae, though sometimes formerly placed in Cephalotaxaceae. Four species are native to eastern Asia; the other two are native to North America. They are small to medium-sized evergreen trees reaching 5–20 m, rarely 25 m, tall. Common names include nutmeg yew.

<span class="mw-page-title-main">Welwitschiaceae</span> Family of plants

Welwitschiaceae is a family of plants of the order Gnetales with one living species, Welwitschia mirabilis, found in southwestern Africa. Three fossil genera have been recovered from the Crato Formation – late Aptian strata located in the Araripe Basin in northeastern Brazil, with one of these also being known from the early Late Cretaceous (Cenomanian-Turonian) Akrabou Formation of Morocco.

<span class="mw-page-title-main">Trimeniaceae</span> Family of flowering plants

Trimeniaceae is a family of flowering plants recognized by most taxonomists, at least for the past several decades. It is a small family of one genus, Trimenia, with eight known species of woody plants, bearing essential oils. The family is subtropical to tropical and found in Southeast Asia, eastern Australia and on several Pacific Islands.

One of the prime systems of plant taxonomy, the Engler system was devised by Adolf Engler (1844–1930), and is featured in two major taxonomic texts he authored or co-authored. His influence is reflected in the use of the terms "Engler School" and "Engler Era". Engler's starting point was that of Eichler who had been the first to use phylogenetic principles, although Engler himself did not think that he was.

Plant taxonomy is the science that finds, identifies, describes, classifies, and names plants. It is one of the main branches of taxonomy.

The Kubitzki system is a system of plant taxonomy devised by Klaus Kubitzki, and is the product of an ongoing survey of vascular plants, entitled The Families and Genera of Vascular Plants, and extending to 15 volumes in 2018. The survey, in the form of an encyclopedia, is important as a comprehensive, multivolume treatment of the vascular plants, with keys to and descriptions of all families and genera, mostly by specialists in those groups. The Kubitzki system served as the basis for classification in Mabberley's Plant-Book, a dictionary of the vascular plants. Mabberley states, in his Introduction on page xi of the 2008 edition, that the Kubitzki system "has remained the standard to which other literature is compared".

<span class="mw-page-title-main">Leptosporangiate fern</span> Subclass of ferns

The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of ferns, the largest of these being the largest group of living ferns, including some 11,000 species worldwide. The group has also been treated as the class Pteridopsida or Polypodiopsida, although other classifications assign them a different rank. Older names for the group include Filicidae and Filicales, although at least the "water ferns" were then treated separately.

<span class="mw-page-title-main">Seed plant</span> Clade of seed plants

A seed plant or spermatophyte, also known as a phanerogam or a phaenogam, is any plant that produces seeds. It is a category of embryophyte that includes most of the familiar land plants, including the flowering plants and the gymnosperms, but not ferns, mosses, or algae.

<span class="mw-page-title-main">Ophioglossidae</span> Subclass of ferns

Ophioglossidae is one of the four subclasses of Polypodiopsida (ferns). This subclass consists of the ferns commonly known as whisk ferns, grape ferns, adder's-tongues and moonworts. It is equivalent to the class Psilotopsida in previous treatments, including Smith et al. (2006). The subclass contains two orders, Psilotales and Ophioglossales, whose relationship was only confirmed by molecular phylogenetic studies.

<span class="mw-page-title-main">Pinidae</span> Subclass of plants

Pinidae is a subclass of Equisetopsida in the sense used by Mark W. Chase and James L. Reveal in their 2009 article "A phylogenetic classification of the land plants to accompany APG III." This subclass comprises the conifers. The Pinidae subclass is equivalent to the division Pinophyta and class Pinopsida of previous treatments. There are over 600 species of Pinidae all over the world.

References

  1. 1 2 Gorozhankin 1904.
  2. 1 2 3 4 5 6 7 8 Christenhusz et al 2011.
  3. 1 2 3 Eckenwalder 2008.
  4. 1 2 Gernandt et al 2011.
  5. Brown 1826.
  6. Tomlinson 2012.
  7. Brongniart 1828–1837.
  8. Lindley 1830.
  9. 1 2 3 4 De-Zhi et al 2004.
  10. Lindley 1853.
  11. Bentham & Hooker 1862–1883.
  12. Engler 1903.
  13. 1 2 Brunsfield et al 1994.
  14. 1 2 Pilger 1926.
  15. Chamberlain 1935.
  16. 1 2 Benson 1957.
  17. Cronquist 1960.
  18. 1 2 Cronquist et al 1966.
  19. 1 2 3 Chase & Reveal 2009.
  20. Farjon 2001.
  21. 1 2 Nagalingum & Burleigh 2018.
  22. Eckenwalder 2009.
  23. Farjon 2010.
  24. Debreczy & Rácz 2011.
  25. Farjon & Filer 2013.
  26. Farjon 2018.
  27. 1 2 3 4 Wang & Ran 2014.
  28. Reveal 1995.
  29. Delevoryas 2015.
  30. 1 2 Cole & Bachelier 2019.
  31. 1 2 3 Lu et al 2014.
  32. Ran et al 2018.
  33. 1 2 Stevens 2020.
  34. Davy 1937.
  35. 1 2 Earle 2020.

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