Conifer

Conifers (/ˈkɒnɪfər/) are a group of seed plants, a subset of gymnosperms. They are mainly evergreen trees with a regular branching pattern, reproducing with male and female cones, usually on the same tree. They are wind-pollinated and the seeds are usually dispersed by the wind. Scientifically, they make up the division Pinophyta, also known as Coniferae. All extant conifers except for the Gnetophytes are perennial woody plants with secondary growth. There are over 600 living species.

Conifers first appear in the fossil record over 300 million years ago in the Carboniferous. They became dominant land plants in the Mesozoic, until flowering plants took over many ecosystems in the Cretaceous. Many conifers today are relict species, surviving in a small part of their former ranges. Such relicts include Wollemia, known only from a small area of Australia, and Metasequoia glyptostroboides, known from Cretaceous fossils and surviving in a small area of China.

Although the total number of species is relatively small, conifers are ecologically important. They are the dominant plants over the taiga of the Northern Hemisphere. Boreal conifers have multiple adaptations to survive winters, including a conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss. Several fungi form ectomycorrhizal associations with conifers. Other fungi cause diseases such as needle cast, especially harmful to young trees. Conifers are affected by pest insects such as wood-boring longhorn beetles and by bark beetles, which make galleries just under the bark. Conifers are of great economic value for timber and paper production.

The earliest conifers appear in the fossil record during the Late Carboniferous (Pennsylvanian) over 300 million years ago. Conifers are thought to be most closely related to the Cordaitales, a group of extinct Carboniferous-Permian trees and clambering plants whose reproductive structures had some similarities to those of conifers. The most primitive conifers belong to the paraphyletic assemblage of "walchian conifers", which were small trees, and probably originated in dry upland habitats. The range of conifers expanded during the Early Permian (Cisuralian) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced voltzialean or "transition" conifers. Conifers were largely unaffected by the Permian–Triassic extinction event, and were dominant land plants of the Mesozoic era. Modern groups of conifers emerged from the Voltziales during the Late Permian through Jurassic. Conifers underwent a major decline in the Late Cretaceous corresponding to the explosive adaptive radiation of flowering plants.

Several extant conifers have relict taxon status, surviving in small areas or in very small numbers where they once may have been common and widespread. One such is Wollemia nobilis, discovered in 1994 in some narrow, steep-sided, sandstone gorges in Australia. The wild population consisted of under 60 adult trees with essentially no genetic variability, implying a genetic bottleneck some thousands of years ago. The extant gnetophytes consist of three relict genera, namely Ephedra, Gnetum, and Welwitschia. Fossils definitely of the group date back to the Late Jurassic, with many species in the Cretaceous. Conifers as a whole, too, declined markedly after the angiosperms (flowering plants) diversified during the Cretaceous, coming to dominate most terrestrial ecosystems. Many conifer species became extinct, leaving 30 out of 80 genera with just one extant species, and 11 more with just two or three species. The popular phrase "living fossils" could, the Dutch botanist Aljos Farjon states, fittingly be applied to many of these. Thus, Metasequoia glyptostroboides, the dawn redwood, is known from fossils of Late Cretaceous and Miocene age, and was found also as an extant tree with a small relict range in China.

The cladogram summarizes the group's external phylogeny. The conifers are gymnosperms, sister to a clade consisting of the ginkgos and cycads.

The Gnetophyta, despite their distinct appearances, were long viewed as outside the conifer group, but phylogenomic analysis indicates that they are part of the conifer clade, sister to the pine family (the 'gnepine' hypothesis). If so, the gnetophytes once shared the distinctive characters of the conifers, and have lost them. The cladogram summarizes the conifers' internal phylogeny:

The name conifer, meaning 'cone-bearing', derives from Latin laconus (cone) and ferre (to bear). As recently as 1999, the botanist Aljos Farjon wrote that while the Coniferae had up to the early 20th century been considered "a natural family", comparable to the Rosaceae, he doubted that the conifers or the gymnosperms formed natural groups (clades). By 2016, the conifers were recognised as a clade, with six families (not including the gnetophytes), 65–70 genera, and over 600 living species (c. 2002). Depending on interpretation, the Cephalotaxaceae may or may not be included within the Taxaceae, while some authors recognize Phyllocladaceae as distinct from Podocarpaceae. The family Taxodiaceae is here included in the family Cupressaceae.