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''Quercus'' subgenus ''Quercus'' is one of the two
subgenera In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
into which the genus ''
Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
'' was divided in a 2017 classification (the other being subgenus ''Cerris''). It contains about 190 species divided among five sections. It may be called the New World clade or the high-latitude clade; most species are native to the Americas, the others being found in
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
and northernmost
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
.


Description

Members of subgenus ''Quercus'' are distinguished from members of subgenus ''Cerris'' by few morphological features, their separation being largely determined by
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
evidence. All are trees or shrubs bearing
acorn The acorn is the nut (fruit), nut of the oaks and their close relatives (genera ''Quercus'', ''Notholithocarpus'' and ''Lithocarpus'', in the family Fagaceae). It usually contains a seedling surrounded by two cotyledons (seedling leaves), en ...
-like fruit in which a cup covers at least the base of the nut. The outer structure of the mature
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
is one feature that distinguishes the two subgenera: in subgenus ''Quercus'', the small folds or wrinkles (rugulae) are obscured by sporopollenin, whereas in subgenus ''Cerris'', the rugulae are visible or at most weakly obscured. The two subgenera are also distinguished to some extent by their different distributions. Subgenus ''Quercus'' occurs mainly in the Americas, with some species native to
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
and
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
, and may be called the New World clade or the high-latitude clade. Subgenus ''Cerris'' is primarily Eurasian, with a few species in North Africa, and may be called the Old World clade or the mid-latitude clade.


Taxonomy


Phylogeny

The following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
summarizes the relationships that Denk et al. used to draw up their 2017 classification:


Section ''Lobatae''

''Quercus'' sect. ''Lobatae'' was established by
John Claudius Loudon John Claudius Loudon (8 April 1782 – 14 December 1843) was a Scottish botanist, garden designer and author, born in Cambuslang in 1782. He was the first to use the term arboretum in writing to refer to a garden of plants, especially trees, co ...
in 1830. The section, or part of it, has also been treated under names including ''Quercus'' sect. ''Rubrae'' Loudon and ''Quercus'' sect. ''Erythrobalanus'' Spach. It has also been treated as the subgenus ''Erythrobalanus'' and as the full genus ''Erythrobalanus'' (Spach) O.Schwarz. Its members may be called red oaks. The
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...
of the pistillate flowers has a characteristic flange-like shape. The staminate flowers have up to six
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s. The stalk connecting the perianth to the
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
is cone-shaped and often has rings. The acorns mature in two years, rarely in one year. The 'cup' (cupule) of the acorn is fused with its stalk ( peduncule) forming a connective piece. Both the connective piece and the cup are covered with small triangular scales, mostly thin and membranous with broadly angled tips. The leaves typically have teeth with bristle-like extensions, or just bristles in leaves without teeth. The section contains about 125 species native to
Northern America Northern America is the northernmost subregion of North America, as well as the northernmost region in the Americas. The boundaries may be drawn significantly differently depending on the source of the definition. In one definition, it lies dir ...
(including
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
),
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
, and
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
in South America. The red oaks of Mexico are one of the groups of oaks that have most rapidly diversified into different species. Molecular evidence suggests that there are significant numbers of undescribed ''Quercus'' species in Mexico, so the number of known species in the section is likely to be an underestimate of the total diversity. File:028 quercus castanea.jpg, '' Quercus castanea'' leaves and acorns File:Quercus coccinea (scarlet oak) 1.jpg, '' Quercus coccinea'' File:Northern Pin Oak.jpg, Leaves of '' Quercus ellipsoidalis'' File:Quercus myrtifolia mosbo6.jpg, '' Quercus myrtifolia'' growing as a shrub File:Red Oak, Prokocim, Krakow, Poland.jpg, ''
Quercus rubra ''Quercus rubra'', the northern red oak, is an oak tree in the red oak group (''Quercus'' section ''Lobatae''). It is a native of North America, in the eastern and central United States and southeast and south-central Canada. It has been intro ...
'' in autumn (in cultivation)


Section ''Protobalanus''

''Quercus'' sect. ''Protobalanus'' was first established as a subgenus by William Trelease in 1922 and then later treated as a section by Otto Karl Anton Schwarz in 1936 and Aimée Antoinette Camus in 1938. The staminate flowers have eight to ten
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s. The
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
ornamentation has only small wrinkles or folds (verrucae). The acorns mature after two years. The cup (cupule) at the base of the acorn has triangular scales that are fused at the base and have sharp angled tips. The scales are thick and compressed into rings, often forming small bumps, that may be obscured by glandular hairs. The leaf teeth end in spines. The section contains only five species, native to southwestern North America and northwestern Mexico. File:Quercus cedrosensis 53513896.jpg, '' Quercus cedrosensis'' in habitat in Mexico File:Quercus chrysolepis 08638.JPG, '' Quercus chrysolepis'' leaves showing spines File:Quercus chrysolepis acorns.jpg, ''Quercus chrysolepis'' acorns File:Island Oak in Santa Rosa Island.jpg, '' Quercus tomentella'' in habitat on Santa Rosa Island, California File:Quercus tomentella acorn.JPG, ''Quercus tomentella'' acorn


Section ''Ponticae''

''Quercus'' sect. ''Ponticae'' was first established by Boris Stefanoff in 1930. It has also been treated as a subsection and a series, including under the name ''Q.'' ser. ''Sadlerianae'' Trelease. Species are shrubs or small trees, with
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s. The staminate
catkin A catkin or ament is a slim, cylindrical flower cluster (a spike), with inconspicuous or no petals, usually wind- pollinated ( anemophilous) but sometimes insect-pollinated (as in '' Salix''). It contains many, usually unisexual flowers, arra ...
s are up to 10 cm long. The cup (cupule) at the base of the acorn has scales with sharp angled ends. The leaves are either evergreen or deciduous, with simple or compound teeth. The leaf buds are large, enclosed in loosely attached scales. There are only two species, '' Quercus pontica'' and '' Quercus sadleriana''. They have disjoint distributions. ''Quercus pontica'' is native to mountainous areas of north-eastern
Turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
and western
Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
. ''Quercus sadleriana'' is native to northern-most
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
and southern-most
Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...
in the United States. File:Quercus pontica-5.jpg, '' Quercus pontica'' in habitat File:Flickr - brewbooks - Quercus sadleriana (Deer Oak).jpg, '' Quercus sadleriana'' in cultivation


Section ''Virentes''

''Quercus'' sect. ''Virentes'' was first established by
John Claudius Loudon John Claudius Loudon (8 April 1782 – 14 December 1843) was a Scottish botanist, garden designer and author, born in Cambuslang in 1782. He was the first to use the term arboretum in writing to refer to a garden of plants, especially trees, co ...
in 1838. It has also been treated as a series. Members of the section may be called live oaks. Species are trees or rhizomatous shrubs. They are
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
or brevideciduous. The acorns mature in a year. The cup at the base of the acorn has narrowly triangular scales, with thin keels, at most small bumps (tubercules), and sharp angled ends. The leaves are evergreen or almost so. A distinctive feature of the section is that the germinating seed has fused seed leaves (
cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...
s) and an elongated stem above the cotyledons (the
epicotyl An epicotyl is important for the beginning stages of a plant's life. It is the region of a seedling stem above the stalks of the seed leaves of an embryo plant. It grows rapidly, showing hypogeal germination, and extends the stem above the soil su ...
) that forms a tube, while the stem below the cotyledons (the
hypocotyl The hypocotyl (short for "hypocotyledonous stem", meaning "below seed leaf") is the stem of a germinating seedling, found below the cotyledons (seed leaves) and above the radicle (root). Eudicots As the plant embryo grows at germination, it send ...
) is
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
ous. The section contains seven species, native to south-eastern Northern America, Mexico, the West Indies (
Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...
), and Central America.


Section ''Quercus''

''Quercus'' sect. ''Quercus'' has been known, either in whole or part, by a variety of names in the past, including ''Quercus'' sect. ''Albae'', ''Quercus'' sect. ''Macrocarpae'' and ''Quercus'' sect. ''Mesobalanus''. Members of the section may be called white oaks. The section includes all white oaks from North America (treated by Trelease as subgenus ''Leucobalanus''). The staminate flowers have seven or more
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s. The
acorn The acorn is the nut (fruit), nut of the oaks and their close relatives (genera ''Quercus'', ''Notholithocarpus'' and ''Lithocarpus'', in the family Fagaceae). It usually contains a seedling surrounded by two cotyledons (seedling leaves), en ...
s mature in one year. The seed leaves (
cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...
s) are either free or fused together. The cup at the base of the acorn has thickened triangular scales that are either free or fused at the base and have sharp angled tips. The scales have keels and are often covered with small bumps (tuberculate). The leaf teeth typically do not have either bristle-like or spiny tips. There are about 150 species, native to Northern America, Mexico, Central America, western
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
, East Asia, and North Africa. File:Keeler Oak Tree - distance photo, May 2013.jpg, Large ''
Quercus alba ''Quercus alba'', the white oak, is one of the preeminent hardwoods of eastern and central North America. It is a long-lived oak, native to eastern and central North America and found from Minnesota, Ontario, Quebec, and southern Maine south as ...
'' growing in
New Jersey New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
File:2021-04-22 18 54 32 Male flowers (catkins) on a White Oak within a wooded area in the Franklin Farm section of Oak Hill, Fairfax County, Virginia.jpg, ''Quercus alba'' catkins (staminate or 'male' flowers) File:Leaves of California Scrub Oak.JPG, '' Quercus berberidifolia'' in habitat in
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
File:Quercus lusitanica - Köhler–s Medizinal-Pflanzen-253.jpg, '' Q. lusitanica'' flowers, staminate (left) and pistillate (top right) File:Chestnut Oak in Weiser State Forest.JPG, '' Q. montana'' in
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
File:Wintereik Engeland.jpg, '' Q. petraea'' in England, about 300 years old File:Летен дъб - цъфтеж, нови листа.JPG, '' Q. robur'', new leaves and pistillate flowers File:Chêne glands.jpg, ''Q. robur'' leaves and acorns


Notes


References

{{Taxonbar, from=Q4163388 Plant subgenera subg. Quercus