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''Tsuga'' (, from Japanese (), the name of ''
Tsuga sieboldii ''Tsuga sieboldii'', also called the southern Japanese hemlock, or in Japanese, simply tsuga (栂), is a conifer native to the Japanese islands of Honshū, Kyūshū, Shikoku and Yakushima. In Europe and North America the tree is sometimes used a ...
'') is a genus of
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s in the subfamily Abietoideae of Pinaceae, the pine family. The
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
hemlock is derived from a perceived similarity in the smell of its crushed foliage to that of the unrelated plant poison hemlock. Unlike the latter, ''Tsuga'' species are not poisonous. The genus comprises eight to ten species (depending on the
authority In the fields of sociology and political science, authority is the legitimate power of a person or group over other people. In a civil state, ''authority'' is practiced in ways such a judicial branch or an executive branch of government.''The N ...
), with four species occurring in North America and four to six in eastern Asia.


Description

They are medium-sized to large evergreen trees, ranging from tall, with a conical to irregular crown, the latter occurring especially in some of the Asian species. The leading shoots generally droop. The
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
is scaly and commonly deeply furrowed, with the colour ranging from grey to brown. The branches stem horizontally from the trunk and are usually arranged in flattened sprays that bend downward towards their tips. Short spur shoots, which are present in many gymnosperms, are weakly to moderately developed. The young twigs, as well as the distal portions of stem, are flexible and often pendent. The stems are rough due to pulvini that persist after the
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
fall. The winter buds are ovoid or globose, usually rounded at the apex and not resinous. The leaves are flattened to slightly angular and range from long and broad. They are borne singly and are arranged spirally on the stem; the leaf bases are twisted so the leaves lie flat either side of the stem or more rarely radially. Towards the base, the leaves narrow abruptly to a petiole set on a forward-angled pulvinus. The petiole is twisted at the base so it is almost parallel with the stem. The leaf apex is either notched, rounded, or acute. The undersides have two white
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
tal bands (which are inconspicuous on '' T. mertensiana'') separated by an elevated midvein. The upper surface of the leaves lack stomata, except those of ''T. mertensiana''. They have one resin canal that is present beneath the single vascular bundle. The
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
cones grow solitary from lateral buds. They are usually up to in length, ovoid, globose, or ellipsoid, and yellowish-white to pale purple, and borne on a short peduncle. The pollen itself has a saccate, ring-like structure at its distal pole, and rarely this structure can be more or less doubly saccate. The seed cones are borne on year-old twigs and are small ovoid-globose or oblong-cylindric, ranging from long, except in ''T. mertensiana'', where they are cylindrical and longer, in length; they are solitary, terminal or rarely lateral, pendulous, and are sessile or on a short peduncle up to long. Maturation occurs in 5–8 months, and the seeds are shed shortly thereafter; the cones are shed soon after seed release or up to a year or two later. The seed scales are thin, leathery, and persistent. They vary in shape and lack an apophysis and an umbo. The
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
s are included and small. The seeds are small, from long, and winged, with the wing being in length. They also contain small adaxial resin
vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
. Seed germination is
epigeal Epigeal, epigean, epigeic and epigeous are biological terms describing an organism's activity above the soil surface. In botany, a seed is described as showing epigeal germination when the cotyledons of the germinating seed expand, throw off the ...
; the seedlings have 4–6 cotyledons.


Taxonomy

Mountain hemlock (''T. mertensiana'') is unusual in the genus in several respects. The leaves are less flattened and arranged all round the shoot, and have stomata above as well as below, giving the foliage a glaucous color; and the cones are the longest in the genus, long and cylindrical rather than ovoid. Some botanists treat it in a distinct genus as ''Hesperopeuce mertensiana'' (Bong.) Rydb., though it is more generally only considered distinct at the rank of subgenus. Another species, bristlecone hemlock, first described as ''T. longibracteata'', is now treated in a distinct genus ''
Nothotsuga ''Nothotsuga'' is a genus of coniferous trees in the family Pinaceae endemic to China. ''Nothotsuga'' contains only one species, ''Nothotsuga longibracteata'', commonly known as the bristlecone hemlock, which is found in southeastern China, in ...
''; it differs from ''Tsuga'' in the erect (not pendulous) cones with exserted bracts, and male cones clustered in umbels, in these features more closely allied to the genus '' Keteleeria''.


Species

; Accepted living species * '' Tsuga canadensis'' – eastern hemlock – Eastern Canada, Eastern United States * '' Tsuga caroliniana'' – Carolina hemlock – Southern Appalachians * '' Tsuga chinensis'' – Taiwan hemlock – much of China incl Tibet + Taiwan * ''
Tsuga diversifolia ''Tsuga diversifolia'', commonly known as the northern Japanese hemlock, or in Japanese, kometsuga (米栂), is a species of conifer native to the Japanese islands of Honshū, Kyūshū, and Shikoku. In Europe and North America, the species is so ...
'' – northern Japanese hemlock – Honshu,
Kyushu is the third-largest island of Japan's five main islands and the most southerly of the four largest islands ( i.e. excluding Okinawa). In the past, it has been known as , and . The historical regional name referred to Kyushu and its surroun ...
* ''
Tsuga dumosa ''Tsuga dumosa'', commonly called the Himalayan hemlock or in Chinese, Yunnan tieshan (), is a species of conifer native to the eastern Himalayas. It occurs in parts of Nepal, India, Bhutan, Myanmar, Vietnam and Tibet. Within its native range t ...
'' – Himalayan hemlock – Himalayas, Tibet, Yunnan, Sichuan * ''
Tsuga forrestii ''Tsuga forrestii'' is a species of conifer in the family Pinaceae. It is considered as a variety of ''Tsuga chinensis'' (i.e., ''T. c.'' var. ''forrestii'') by some. It grows in mixed forests in mountains and valleys of northeast Guizhou, south ...
'' – Forrest's hemlock – Sichuan, Yunnan, Guizhou * '' Tsuga heterophylla'' – western hemlock – Western Canada, Northwestern United States * ''
Tsuga × jeffreyi ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed folia ...
'' – British Columbia, Washington * '' Tsuga mertensiana'' – mountain hemlock – Alaska, British Columbia, Western United States * ''
Tsuga sieboldii ''Tsuga sieboldii'', also called the southern Japanese hemlock, or in Japanese, simply tsuga (栂), is a conifer native to the Japanese islands of Honshū, Kyūshū, Shikoku and Yakushima. In Europe and North America the tree is sometimes used a ...
'' – southern Japanese hemlock – Japan * '' Tsuga ulleungensis'' – Ulleungdo hemlock – Ulleungdo island, Korea ;Accepted paleospecies * †''
Tsuga aburaensis ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed folia ...
'' - Abura, Hokkaido (Miocene) * †''
Tsuga asiatica ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed foliage ...
'' - Lawula Formation, Tibet (
Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage of t ...
) * †''
Tsuga europaea ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed foliage ...
'' -Maria Theresia mine
Aachen Aachen ( ; ; Aachen dialect: ''Oche'' ; French and traditional English: Aix-la-Chapelle; or ''Aquisgranum''; nl, Aken ; Polish: Akwizgran) is, with around 249,000 inhabitants, the 13th-largest city in North Rhine-Westphalia, and the 28th- ...
(Miocene) * †''
Tsuga nanfengensis ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed foliage ...
'' - Yunnan Province (Late Miocene) * †'' Tsuga swedaea'' -
Buchanan Lake Formation The Buchanan Lake Formation is a geologic formation on Axel Heiberg Island in Nunavut, and part of the Eureka Sound Group. The formation is divided into four members and preserves fossils dating back to the Lutetian stage. See also * List of ...
,
Axel Heiberg Island Axel Heiberg Island ( iu, ᐅᒥᖕᒪᑦ ᓄᓈᑦ, ) is an uninhabited island in the Qikiqtaaluk Region, Nunavut, Canada. Located in the Arctic Ocean, it is the 32nd largest island in the world and Canada's seventh largest island. According ...
( Lutetian) * †''
Tsuga taxoides ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed foliage ...
'' - Inner Mongolia (Early Cretaceous) * †''
Tsuga xianfengensis ''Tsuga'' (, from Japanese (), the name of ''Tsuga sieboldii'') is a genus of conifers in the subfamily Abietoideae of Pinaceae, the pine family. The common name hemlock is derived from a perceived similarity in the smell of its crushed folia ...
'' - Yunnan Province (late Miocene) ; Formerly included Moved to other genera: ''
Cathaya ''Cathaya'' is a genus in the pine family, Pinaceae, with one known living species, ''Cathaya argyrophylla''. ''Cathaya'' is a member of the subfamily Laricoideae, most closely related to ''Pseudotsuga'' and ''Larix''. A second species, ''C. nan ...
Keteleeria
Nothotsuga ''Nothotsuga'' is a genus of coniferous trees in the family Pinaceae endemic to China. ''Nothotsuga'' contains only one species, ''Nothotsuga longibracteata'', commonly known as the bristlecone hemlock, which is found in southeastern China, in ...
Picea A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
Pseudotsuga ''Pseudotsuga'' is a genus of evergreen coniferous trees in the family Pinaceae (subfamily Laricoideae). Common names for species in the genus include Douglas fir, Douglas-fir, Douglas tree, Oregon pine and Bigcone spruce. '' Pseudotsuga menz ...
Taxus''


Ecology

The species are all adapted to (and are confined to) relatively moist, cool temperate areas with high rainfall, cool summers, and little or no water stress; they are also adapted to cope with heavy to very heavy winter snowfall and tolerate ice storms better than most other trees. Hemlock trees are more tolerant of heavy shade than other conifers; they are, however, more susceptible to drought.


Threats

The two eastern North American species, '' T. canadensis'' and '' T. caroliniana'', are under serious threat by the sap-sucking insect ''Adelges tsugae'' ( hemlock woolly adelgid). This adelgid, related to the aphids, was introduced accidentally from eastern Asia, where it is only a minor pest. Extensive mortality has occurred, particularly east of the Appalachian Mountains. The Asian species are resistant to this pest, and the two western American hemlocks are moderately resistant. In North America, hemlocks are also attacked by
hemlock looper ''Lambdina fiscellaria'', the mournful thorn or hemlock looper, is a moth of the family Geometridae. It is found in North America, from the Pacific to the Atlantic coast and from Canada south to Pennsylvania, Wisconsin and California. The adult ...
.USDA Forest Service NA-PR-05-92 Larger infected hemlocks have large, relatively high root systems that can bring other trees down if one falls. The foliage of young trees is often browsed by deer, and the seeds are eaten by finches and small rodents. Old trees are commonly attacked by various fungal disease and decay species, notably ''
Heterobasidion annosum ''Heterobasidion annosum'' is a basidiomycete fungus in the family Bondarzewiaceae. It is considered to be the most economically important forest pathogen in the Northern Hemisphere. ''Heterobasidion annosum'' is widespread in forests in the Unit ...
'' and '' Armillaria'' species, which rot the heartwood and eventually leave the tree liable to windthrow, and ''
Rhizina undulata ''Rhizina undulata'', commonly known as the doughnut fungus or the pine firefungus, is a species of fungus in the family Rhizinaceae. The fruit bodies of the fungus are dark purple brown with a bright yellow margin, crust-like and attached to the ...
'', which may kill groups of trees following minor grass fires that activate growth of the ''Rhizina'' spores.Phillips, D. H., & Burdekin, D. A. (1992). ''Diseases of Forest and Ornamental Trees''. Macmillan .


Uses

The wood obtained from hemlocks is important in the timber industry, especially for use as wood pulp. Many species are used in horticulture, and numerous cultivars have been selected for use in gardens. The bark of the hemlock is also used in tanning leather. The needles of the hemlock tree are sometimes used to make a tea and perfume.


References


External links

* * {{Authority control Conifer genera