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Sequoioideae, popularly known as redwoods, is a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
of
coniferous 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 ...
trees within the family
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdio ...
. It includes the largest and tallest trees in the world.


Description

The three redwood subfamily
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
are '' Sequoia'' from coastal California and Oregon, '' Sequoiadendron'' from California's
Sierra Nevada The Sierra Nevada () is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primarily ...
, and '' Metasequoia'' in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. The redwood species contains the largest and tallest trees in the world. These trees can live for thousands of years. Threats include logging, fire suppression, climate change, illegal marijuana cultivation, and burl poaching. Only two of the genera, ''Sequoia'' and ''Sequoiadendron'', are known for massive trees. Trees of '' Metasequoia'', from the single living species '' Metasequoia glyptostroboides'', are much smaller.


Taxonomy and evolution

Multiple studies of both morphological and molecular characters have strongly supported the assertion that the Sequoioideae are
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. Most modern phylogenies place ''Sequoia'' as sister to ''Sequoiadendron'' and ''Metasequoia ''as the out-group. However, Yang et al. went on to investigate the origin of a peculiar genetic artifact of the Sequoioideae—the polyploidy of ''Sequoia''—and generated a notable exception that calls into question the specifics of this relative consensus.


Cladistic tree

A 2006 paper based on non-molecular evidence suggested the following relationship among extant species: A 2021 study using molecular evidence found the same relationships among Sequoioideae species, but found Sequoioideae to be the sister group to the
Athrotaxidoideae ''Athrotaxis'' is a genus of two to three species (depending on taxonomic opinion) of conifers in the cypress family, Cupressaceae. The genus is endemic to western Tasmania, where they grow in high altitude temperate rainforests.Farjon, A. (200 ...
(a superfamily presently known only from Tasmania) rather than to Taxodioideae. Sequoioideae and Athrotaxidoideae are thought to have diverged from each other during the Jurassic.


Possible reticulate evolution in Sequoioideae

Reticulate evolution refers to the origination of a taxon through the merging of ancestor lineages. Polyploidy has come to be understood as quite common in plants—with estimates ranging from 47% to 100% of flowering plants and extant ferns having derived from ancient polyploidy. Within the gymnosperms however it is quite rare. ''Sequoia sempervirens'' is hexaploid (2n= 6x= 66). To investigate the origins of this polyploidy Yang et al. used two single copy nuclear genes, LFY and NLY, to generate
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
s. Other researchers have had success with these genes in similar studies on different taxa. Several hypotheses have been proposed to explain the origin of ''Sequoia's ''polyploidy: allopolyploidy by hybridization between ''Metasequoia'' and some probably extinct taxodiaceous plant; ''Metasequoia'' and ''Sequoiadendron,'' or ancestors of the two genera, as the parental species of ''Sequoia''; and autohexaploidy, autoallohexaploidy, or segmental allohexaploidy. Yang et al. found that ''Sequoia'' was clustered with ''Metasequoia'' in the tree generated using the LFY gene but with ''Sequoiadendron ''in the tree generated with the NLY gene. Further analysis strongly supported the hypothesis that ''Sequoia'' was the result of a hybridization event involving ''Metasequoia'' and ''Sequoiadendron''. Thus, Yang et al. hypothesize that the inconsistent relationships among ''Metasequoia'', ''Sequoia,'' and ''Sequoiadendron'' could be a sign of reticulate evolution by hybrid speciation (in which two species hybridize and give rise to a third) among the three genera. However, the long evolutionary history of the three genera (the earliest fossil remains being from the Jurassic) make resolving the specifics of when and how ''Sequoia'' originated once and for all a difficult matter—especially since it in part depends on an incomplete fossil record.


Extant Species


South-central China


California, USA


Paleontology

Sequoioideae is an ancient taxon, with the oldest described Sequoioideae species, ''Sequoia jeholensis'', recovered from Jurassic deposits.Ahuja M. R. and D. B. Neale. 2002
Origins of polyploidy in coast redwood (Sequoia sempervirens) and relationship of coast redwood (Sequoia sempervirens) to other genera of Taxodiaceae.
Silvae Genetica 51: 93–99.
A genus ''Medulloprotaxodioxylon'', reported from the late Triassic of China supports the idea of a
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
origin. The fossil record shows a massive expansion of range in the Cretaceous and dominance of the Arcto-Tertiary Geoflora, especially in northern latitudes. Genera of Sequoioideae were found in the
Arctic Circle The Arctic Circle is one of the two polar circles, and the most northerly of the five major circles of latitude as shown on maps of Earth. Its southern equivalent is the Antarctic Circle. The Arctic Circle marks the southernmost latitude at w ...
, Europe, North America, and throughout Asia and Japan. A general cooling trend beginning in the late Eocene and
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
reduced the northern ranges of the Sequoioideae, as did subsequent ice ages. Evolutionary adaptations to ancient environments persist in all three species despite changing climate, distribution, and associated flora, especially the specific demands of their reproduction ecology that ultimately forced each of the species into refugial ranges where they could survive.


Conservation

The entire subfamily is endangered. The IUCN Red List Category & Criteria assesses ''Sequoia sempervirens'' as Endangered (A2acd), ''Sequoiadendron giganteum'' as Endangered (B2ab) and ''Metasequoia glyptostroboides'' as Endangered (B1ab).


Introduced range

The two California redwood species, since the early 19th century, and the Chinese redwood species since 1948, have been cultivated horticulturally far beyond their native habitats. They are found in botanical gardens, public parks, and private landscapes in many similar climates worldwide. Plantings outside their native ranges particularly are found in California, the coastal Northwestern and the Eastern United States, areas of China, Ireland, Germany, the United Kingdom, Australia and near
Rotorua Rotorua () is a city in the Bay of Plenty region of New Zealand's North Island. The city lies on the southern shores of Lake Rotorua, from which it takes its name. It is the seat of the Rotorua Lakes District, a territorial authority encompass ...
, New Zealand. They are also used in educational projects recreating the look of the megaflora of the Pleistocene landscape.


See also

* Temperate
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...
of North America Westcoast (Sequoia forests) * List of superlative trees


References


Bibliography and links

* * * * * :de:Liste der dicksten Mammutbäume in Deutschland. List of Large Giant Redwoods in Germany * IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. Downloaded on 10 January 2014. * *


External links

* {{Authority control Plant subfamilies