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''Torreya'' is a genus of
conifers 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 ext ...
comprising six or seven species placed in the family
Taxaceae Taxaceae (), commonly called the yew family, is a coniferous family which includes six extant and two extinct genera, and about 30 species of plants, or in older interpretations three genera and 7 to 12 species. Description They are many-bran ...
, though sometimes formerly placed in
Cephalotaxaceae Cephalotaxaceae is a small grouping of conifers, that included one to three genera closely allied to Taxaceae. However, members of Cephalotaxaceae are now included in Taxaceae by botanists, instead of as a distinct family, based on phylogenetic e ...
. Four species are native to eastern
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
; the other two are native to
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
. They are small to medium-sized
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, whic ...
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s reaching 5–20 m, rarely 25 m, tall. Common names include nutmeg yew. The genus is one example 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 18 ...
in
paleoecology Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
. The pattern of highly
disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
of geographic ranges of the species within such a genus spans temperate plant zones of continents in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
. This geographic pattern is attributed to genus origins in much warmer times of the
Tertiary Period Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
, when zones of
temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
were found in poleward latitudes whereby land connections facilitated range expansions and migrations of plants between Asia and North America and sometimes between Europe and North America. The
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
are spirally arranged on the shoots, but twisted at the base to lie in two flat ranks; they are linear, 2–8 cm long and 3–4 mm broad, hard in texture, with a sharp spine tip. ''Torreya'' can be
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is ...
,
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...
, or subdioecious (documented in Japanese Torreya and Florida Torreya). When monoecious, the male and female cones are often on different branches. The male (
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 gametop ...
) cones are 5–8 mm long, grouped in lines along the underside of a shoot. The female (
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
) cones are single or grouped two to eight together on a short stem; minute at first, they mature in about 18 months to a
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part ( exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or ''pyrena'') of hardened endocarp with a seed (''kernel ...
-like structure with the single large nut-like seed 2–4 cm long surrounded by a fleshy covering, green to purple at full maturity. In some species, notably the
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the n ...
ese '' Torreya nucifera'' ('kaya'), and unusually for members of Taxaceae, the seed is edible. Natural dispersal is thought to be aided by
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. ...
s which bury the seeds for a winter food source; any seeds left uneaten are then able to germinate. The genus is named after the American botanist
John Torrey John Torrey (August 15, 1796 – March 10, 1873) was an American botanist, chemist, and physician. Throughout much of his career, he was a teacher of chemistry, often at multiple universities, while he also pursued botanical work, focusing on t ...
.


Fossil record

†'' Torreya clarnensis'' has been described from
Middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "da ...
fossils found in the
Clarno Formation Clarno may refer to: Locations * Clarno, Oregon *Clarno, Wisconsin **Clarno (community), Wisconsin *Clarno Township, Lake County, South Dakota Lake County is a County (United States), county in the U.S. state of South Dakota. As of the 2020 Uni ...
of Central
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 ...
,
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
. Leafy branch
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
of †'' Torreya bilinica'' are known from
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 t ...
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
of Zichov near
Bílina Bílina (; german: Bilin) is a town in Teplice District in the Ústí nad Labem Region of the Czech Republic. It has about 14,000 inhabitants. It is known for its spas and as a source of the strongly mineralized water, Bílinská kyselka. The to ...
,
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
, early
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
deposits of Güvem central
Anatolia Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
,
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a transcontinental country located mainly on the Anatolian Peninsula in Western Asia, with a small portion on the Balkan Peninsula ...
and late
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
deposits of
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = '' Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , ...
. A single ''Torreya nucifera'' needle leaf is known from a late Miocene deposit in
Abkhazia Abkhazia, ka, აფხაზეთი, tr, , xmf, აბჟუა, abzhua, or ( or ), officially the Republic of Abkhazia, is a partially recognised state in the South Caucasus, recognised by most countries as part of Georgia, which ...
. More complete records of ''Torreya nucifera'' are known from
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
. Fossil pollen of genus ''Torreya'' and other genera within
Taxaceae Taxaceae (), commonly called the yew family, is a coniferous family which includes six extant and two extinct genera, and about 30 species of plants, or in older interpretations three genera and 7 to 12 species. Description They are many-bran ...
is generally deemed indistinguishable, one from another, and also from genera within families
Taxodiaceae Taxodiaceae is a formerly recognized coniferous plant family comprising the following ten genera: *''Athrotaxis'' *''Cryptomeria'' *''Cunninghamia'' *†'' Cunninghamites'' *''Glyptostrobus'' *''Metasequoia'' *'' Sciadopitys'' *'' Sequoia'' *''S ...
and
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, subdioeci ...
. Therefore, it is generally difficult to support past presence or absence of such genera in geographic locales where macrofossil plant material is rare or absent, even if substantial pollen (as in
Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS). It follows the Neogene Period and spans from 2.58 million year ...
bogs) is available.


Adaptive growth forms of this subcanopy genus

Species within this genus are all adapted to establish and grow slowly as subcanopy woody plants in forest habitats of moderate to dense shade. Stems will lean in very shady conditions, with branches and additional stems arising from the root crown growing more horizontally than vertically. If and when a canopy opening occurs, upward growth will be stimulated and even the leaf form and orientation will shift. Seed production occurs on female branches (and trees) only in the presence of direct sunlight. Therefore, wild trees tended and planted trees cultivated for local or market seed consumption in Asia (usually, ''Torreya grandis'') will have their surroundings managed accordingly through human intervention.


Species


References


Gymnosperm Database: ''Torreya''

Torreya State Park (Florida)
{{Taxonbar, from=Q429944 Conifer genera