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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 Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
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 California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
and
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
, ''
Sequoiadendron ''Sequoiadendron'' is a genus of evergreen trees, with two species, only one of which survives to the present: * ''Sequoiadendron giganteum'', extant, commonly known as wellingtonia, giant redwood and giant sequoia, growing naturally in the Sie ...
'' 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 ''Metasequoia'', or dawn redwoods, is a genus of fast-growing deciduous trees, one of three species of pinophyta, conifers known as redwoods. The living species ''Metasequoia glyptostroboides'' is native to Lichuan, Hubei, Lichuan county in Hube ...
'' 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 In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
, illegal marijuana cultivation, and
burl A burl (American English) or burr (British English) is a tree growth in which the grain has grown in a deformed manner. It is commonly found in the form of a rounded outgrowth on a tree trunk or branch that is filled with small knots from do ...
poaching. Only two of the genera, ''Sequoia'' and ''Sequoiadendron'', are known for massive trees. Trees of ''
Metasequoia ''Metasequoia'', or dawn redwoods, is a genus of fast-growing deciduous trees, one of three species of pinophyta, conifers known as redwoods. The living species ''Metasequoia glyptostroboides'' is native to Lichuan, Hubei, Lichuan county in Hube ...
'', from the single living species ''
Metasequoia glyptostroboides ''Metasequoia glyptostroboides'', the dawn redwood, is a fast-growing, endangered deciduous pinophyta, conifer. It is the sole living species of the genus ''Metasequoia'', one of three genera in the subfamily Sequoioideae of the family (botany), ...
'', 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 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 ...
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 Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
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 ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
) rather than to
Taxodioideae Taxodioideae is a subfamily in Cupressaceae. Genera See also *Taxodiaceae Taxodiaceae is a formerly recognized coniferous plant family comprising the following ten genera: *''Athrotaxis'' *''Cryptomeria'' *''Cunninghamia'' *†'' Cunningham ...
. Sequoioideae and Athrotaxidoideae are thought to have diverged from each other during the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
.


Possible reticulate evolution in Sequoioideae

Reticulate evolution Reticulate evolution, or network evolution is the origination of a lineage through the partial merging of two ancestor lineages, leading to relationships better described by a phylogenetic network than a bifurcating tree. Reticulate patterns ca ...
refers to the origination of a taxon through the merging of ancestor lineages.
Polyploidy Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
has come to be understood as quite common in plants—with estimates ranging from 47% to 100% of flowering plants and extant
ferns A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except th ...
having derived from ancient polyploidy. Within the gymnosperms however it is quite rare. ''Sequoia sempervirens'' is
hexaploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
(2n= 6x= 66). To investigate the origins of this polyploidy Yang et al. used two single copy
nuclear gene A nuclear gene is a gene whose physical DNA nucleotide sequence is located in the cell nucleus of a eukaryote. The term is used to distinguish nuclear genes from genes found in mitochondria or chloroplasts. The vast majority of genes in eukaryote ...
s, 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 Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
by
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
ization 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 Reticulate evolution, or network evolution is the origination of a lineage through the partial merging of two ancestor lineages, leading to relationships better described by a phylogenetic network than a bifurcating tree. Reticulate patterns ca ...
by
hybrid speciation Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was th ...
(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 The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
) 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 In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, with the oldest described Sequoioideae species, ''Sequoia jeholensis'', recovered from
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
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 The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
and dominance of the
Arcto-Tertiary Geoflora The Arcto-Tertiary Geoflora is a hypothesized floral assemblage that once covered the Northern Hemisphere, from roughly the late Mesozoic to mid Cenozoic Eras. Origins First proposed by the paleobotanists J.S. Gardner and C. Ettinghausen in 1879 ...
, 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 The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''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 An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
. The
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
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 garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...
s, 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 The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
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 The world's superlative trees can be ranked by any factor. Records have been kept for trees with superlative height, trunk diameter or girth, canopy coverage, airspace volume, wood volume, estimated mass, and age. Tallest The heights of th ...


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