The Fagales are an order of

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s, including some of the best-known

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. The order name is derived from

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

''Fagus'',

beech Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...

es. They belong among the

rosid The rosids are members of a large clade ( monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classifica ...

group of dicotyledons. The families and genera currently included are as follows: *

Betulaceae Betulaceae, the birch family, includes six genera of deciduous nut-bearing trees and shrubs, including the birches, alders, hazels, hornbeams, hazel-hornbeam, and hop-hornbeams numbering a total of 167 species. They are mostly natives of the t ...

– birch family (''

Alnus Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...

'', ''

Betula A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 3 ...

'', ''

Carpinus Hornbeams are hardwood trees in the flowering plant genus ''Carpinus'' in the birch family Betulaceae. The 30–40 species occur across much of the temperate regions of the Northern Hemisphere. Origin of names The common English name ''hornbea ...

'', ''

Corylus The hazel (''Corylus'') is a genus of deciduous trees and large shrubs native to the temperate Northern Hemisphere. The genus is usually placed in the birch family Betulaceae,Germplasmgobills Information Network''Corylus''Rushforth, K. (1999). ...

'', ''

Ostrya ''Ostrya'' is a genus of eight to 10 small deciduous trees belonging to the birch family Betulaceae. Common names include hop-hornbeam and hophornbeam. It may also be called ironwood, a name shared with a number of other plants. The genus is na ...

'', ''

Ostryopsis ''Ostryopsis'' is a small genus of deciduous shrubs belonging to the birch family Betulaceae. The species have no common English name, though hazel-hornbeam has been suggested, reflecting their similarities to the closely related hazels and h ...

'') *

Casuarinaceae The Casuarinaceae are a family of dicotyledonous flowering plants placed in the order Fagales, consisting of four genera and 91 species of trees and shrubs native to eastern Africa, Australia, Southeast Asia, Malesia, Papuasia, and the Pacifi ...

– she-oak family (''

Allocasuarina ''Allocasuarina'' is a genus of trees in the flowering plant family Casuarinaceae. They are endemic to Australia, occurring primarily in the south. Like the closely related genus ''Casuarina'', they are commonly called sheoaks or she-oaks. Wi ...

'', ''

Casuarina ''Casuarina'' is a genus of 17 tree species in the family Casuarinaceae, native to Australia, the Indian subcontinent, southeast Asia, islands of the western Pacific Ocean, and eastern Africa. It was once treated as the sole genus in the fami ...

'', ''

Ceuthostoma ''Ceuthostoma'' is a genus of two species of trees, constituting part of the plant family Casuarinaceae. They grow naturally only in Malesia, in the islands of Palawan, Borneo, Halmahera and New Guinea. Species * ''Ceuthostoma palawanense'' ...

'', ''

Gymnostoma ''Gymnostoma'' is a genus of about eighteen species of trees and shrubs, constituting one of the four genera of the plant family Casuarinaceae. The species grow naturally in the tropics, including at high altitudes having temperate climates, in ...

'') *

Fagaceae The Fagaceae are a family of flowering plants that includes beeches, chestnuts and oaks, and comprises eight genera with about 927 species. Fagaceae in temperate regions are mostly deciduous, whereas in the tropics, many species occur as evergre ...

– beech family ('' Castanea'', ''

Castanopsis ''Castanopsis'', commonly called chinquapin or chinkapin, is a genus of evergreen trees belonging to the beech family, Fagaceae. The genus contains about 140 species, which are today restricted to tropical and subtropical eastern Asia. A total o ...

'', ''

Chrysolepis ''Chrysolepis'' is a small genus of plants in the family Fagaceae, endemic to the western United States. Its two species have the common name chinquapin. The genus occurs from western Washington south to the Transverse Ranges in Southern Califo ...

'', ''

Colombobalanus ''Trigonobalanus'' is a genus of three species of evergreen trees in the family Fagaceae, related to oaks, beeches and chestnuts. The species are widely scattered, with one in northern South America and two in southeast Asia; some botanists t ...

'', ''

Fagus Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...

'', '' Lithocarpus'', ''

Notholithocarpus ''Notholithocarpus densiflorus'', commonly known as the tanoak or tanbark-oak, is a broadleaf tree in the family Fagaceae, and the type species of the genus ''Notholithocarpus''. It is native to the far western United States, particularly Oregon ...

'', ''

Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...

'') *

Juglandaceae The Juglandaceae are a plant family known as the walnut family. They are trees, or sometimes shrubs, in the order Fagales. Members of this family are native to the Americas, Eurasia, and Southeast Asia. The nine or ten genera in the family have ...

– walnut family (''

Alfaroa ''Alfaroa'' is a genus of evergreen trees in the Juglandaceae family of the Fagales, growing in montane and submontane tropical rain forests in Central America. The wood is characterized by solid pith, pink heartwood, and vessels with scalarifor ...

'', ''

Carya Hickory is a common name for trees composing the genus ''Carya'', which includes around 18 species. Five or six species are native to China, Indochina, and India (Assam), as many as twelve are native to the United States, four are found in Mexi ...

'', ''

Cyclocarya ''Cyclocarya'' is a genus of flowering plants in the family Juglandaceae, comprising a single species ''Cyclocarya paliurus'' (qing qian liu or wheel wingnut), formerly treated in the genus ''Pterocarya'' as ''Pterocarya paliurus''. It is native ...

'', ''

Engelhardia ''Engelhardia'' is a genus of seven species of trees in the family Juglandaceae, native to southeast Asia from northern India east to Taiwan, Indonesia and the Philippines. The genus name is commonly misspelled "''Engelhardtia''", a "correctio ...

'', ''

Juglans Walnut trees are any species of tree in the plant genus ''Juglans'', the type genus of the family Juglandaceae, the seeds of which are referred to as walnuts. All species are deciduous trees, tall, with pinnate leaves , with 5–25 leaflets ...

'', ''

Oreomunnea ''Oreomunnea'' is a genus of two species of flowering plants in the family Juglandaceae, native to southern Mexico and Central America, where they occur in montane rainforest.Manning, W. E. (1979). The Classification within the Juglandaceae. ''An ...

'', ''

Platycarya ''Platycarya'' is a genus of flowering plants in the family Juglandaceae native to eastern Asia in China, Korea, and Japan.Flora of China''Platycarya''/ref>Flora of China''Platycarya strobilacea''/ref> It is a deciduous tree growing to 15 m ta ...

'', ''

Pterocarya ''Pterocarya'', often called wingnuts in English, are trees in the walnut family Juglandaceae. They are native to Asia. The botanic name is from Ancient Greek (''pteron'') "wing" + (''karyon'') " nut". Description ''Pterocarya'' are deciduous ...

'', ''

Rhoiptelea ''Rhoiptelea'' is a monotypic genus of flowering plants in the family Juglandaceae. It contains a single species, ''Rhoiptelea chiliantha'', commonly known as the horsetail tree. This genus was previously recognized in its own family, Rhoipteleac ...

'') *

Myricaceae The Myricaceae are a small family of dicotyledonous shrubs and small trees in the order Fagales. There are three genera in the family, although some botanists separate many species from ''Myrica'' into a fourth genus ''Morella''. About 55 speci ...

– bayberry family (''

Canacomyrica ''Canacomyrica'' is a monotypic genus of flowering plants in the family Myricaceae containing the single species ''Canacomyrica monticola''. It is endemic to New Caledonia. This endangered tree or small shrub is limited to ultramafic serpentine s ...

'', '' Comptonia'', ''

Myrica ''Myrica'' is a genus of about 35–50 species of small trees and shrubs in the family Myricaceae, order Fagales. The genus has a wide distribution, including Africa, Asia, Europe, North America and South America, and missing only from Austral ...

'') *

Nothofagaceae ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...

– southern beech family (''

Nothofagus ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...

'') *

Ticodendraceae ''Ticodendron incognitum'' is the only species of ''Ticodendron'', and the only member of the family Ticodendraceae. It is most closely related to the family Betulaceae. It was discovered only in 1989 in Costa Rica, having been overlooked previ ...

– ticodendron family ('' Ticodendron'') The older

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) a ...

only included four families (Betulaceae, Corylaceae, Fagaceae, Ticodendraceae; Corylaceae now being included within Betulaceae); this arrangement is followed by, for example, the World Checklist of selected plant families. The other families were split into three different orders, placed among the

Hamamelidae Hamamelididae is an obsolete botanical name at the rank of subclass. Because some hamamelidid members bear aments (''i.e.'', catkins), this subclass has been formerly known as ''Amentiferae''. Based on molecular phylogeny works, Hamamelididae app ...

. The Casuarinales comprised the single family Casuarinaceae, the Juglandales comprised the Juglandaceae and Rhoipteleaceae, and the Myricales comprised the remaining forms (plus ''

Balanops ''Balanops'' is a group of flowering plants described as a genus in 1871. The nine species are trees or shrubs, found in New Caledonia, Fiji, Vanuatu, and northern Queensland. They are dioecious, with separate male and female plants. The genus ...

''). The change is due to studies suggesting the Myricales, so defined, are

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

to the other two groups.

Characteristics

Most Fagales are wind pollinated and are

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 conne ...

with unisexual flowers.

Evolutionary history

The oldest member of the order is the flower '' Soepadmoa cupulata'' preserved in the late

Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...

Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...

New Jersey amber New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of th ...

, which is a

mosaic A mosaic is a pattern or image made of small regular or irregular pieces of colored stone, glass or ceramic, held in place by plaster/mortar, and covering a surface. Mosaics are often used as floor and wall decoration, and were particularly pop ...

with characteristics characteristic of both ''Nothofagus'' and other Fagales, suggesting that the ancestor of all Fagales was ''Nothofagus''-like.

Systematics

Modern

molecular phylogenetics 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 ...

suggest the following relationships:

References

External links

Missouri Botanical Gardens - Fagales
{{Taxonbar, from=Q21881 Angiosperm orders