family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
, commonly known as the rueRUTACEAE in BoDD – Botanical Dermatology Database or
citrus
''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
family, of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s, usually placed in the orderSapindales.
Species of the family generally have
flower
Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s that divide into four or five parts, usually with strong scents. They range in form and size from herbs to
shrub
A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s and large
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, e.g., including only woody plants with secondary growth, only ...
s.
The most economically important
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
in the family is ''
Citrus
''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
'', which includes the orange (''C.'' × ''sinensis''),
lemon
The lemon (''Citrus'' × ''limon'') is a species of small evergreen tree in the ''Citrus'' genus of the flowering plant family Rutaceae. A true lemon is a hybrid of the citron and the bitter orange. Its origins are uncertain, but some ...
(''C.'' × ''limon''), grapefruit (''C.'' × ''paradisi''), and lime (various). '' Boronia'' is a large Australian genus, some members of which are plants with highly fragrant flowers and are used in commercial oil production. Other large genera include ''
Zanthoxylum
''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the Family (biology), family Rutaceae that are native to warm temperate and Subtropics, subtropical areas worldwide. It is the type gen ...
,'' several species of which are cultivated for
Sichuan pepper
Sichuan pepper (, also known as Sichuanese pepper, Szechuan pepper, Chinese prickly ash, Chinese pepper, Mountain pepper, and ''mala'' pepper, is a spice commonly used in Sichuan cuisine in China, Bhutan and in northeast India. It is called mej ...
genera
Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
Most species are trees or shrubs, a few are herbs (the type genus '' Ruta'', '' Boenninghausenia'', and '' Dictamnus''), frequently aromatic with
gland
A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...
s on the
leaves
A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
, sometimes with thorns. The leaves are usually opposed and compound and without
stipule
In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...
s. Pellucid glands, a type of oil gland, are found in the leaves responsible for the aromatic smell of the family's members; traditionally they have been the primary synapomorphic characteristic to identify the Rutaceae.
Flowers are
bract
In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale.
Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
raceme
A raceme () or racemoid is an unbranched, indeterminate growth, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are ...
, and mainly pollinated by insects. They are radially or (rarely) laterally symmetric and generally hermaphroditic. They have four or five—sometimes three—mostly separate
petal
Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s and
sepal
A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106
Etymology
The term ''sepalum'' ...
s and 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 ...
Citrus
''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
''), usually separate or in several groups. Usually they have only a single stigma with 2 to 5 united
carpel
Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...
s. Their ovaries are sometimes separate, but their styles are combined.
The fruit of the Rutaceae are very variable:
berries
A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone fruit, stone or pit (fruit), pit although many wikt:pip#Etymology 2, pips or seeds may be p ...
,
drupe
In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
Sapindaceae
The Sapindaceae are a family (biology), family of flowering plants in the order Sapindales known as the soapberry family. It contains 138 genera and 1,858 accepted species. Examples include Aesculus, horse chestnut, maples, ackee and lychee.
The ...
In 1896, Engler published a division of the family Rutaceae into seven subfamilies. One, Rhabdodendroideae, is no longer considered to belong to the Rutaceae, being treated as the segregate family Rhabdodendraceae, containing only the genus '' Rhabdodendron''. Two monogeneric subfamilies, Dictyolomatoideae and Spathelioideae, are now included in the subfamily Cneoroideae, along with genera Engler placed in other families. The remaining four Engler subfamilies were
Aurantioideae
Aurantioideae (sometimes known as Citroideae) is the subfamily within the rue and citrus family (biology), family (Rutaceae) that contains the citrus. The subfamily's center of diversity is in the monsoon region of eastern Australasia, extending ...
, Rutoideae, Flindersioideae and Toddalioideae. Engler's division into subfamilies largely relied on the characteristics of the fruit, as did others used until
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 ...
methods were applied.
Molecular methods have shown that only Aurantioideae can be clearly differentiated from other members of the family based on fruit. They have not supported the circumscriptions of Engler's three other main subfamilies. In 2012, Groppo et al. divided Rutaceae into only two subfamilies, retaining Cneoroideae but placing all the remaining genera in a greatly enlarged subfamily Rutoideae ''s.l.'' A 2014 classification by Morton and Telmer also retained Engler's Aurantioideae, but split the remaining Rutoideae ''s.l.'' into a smaller Rutoideae and a much larger Amyridoideae ''s.l.'', containing most of Engler's Rutoideae. Until 2021, molecular phylogenetic methods had only sampled between 20% and 40% of the genera of Rutaceae. A 2021 study by Appelhans et al. sampled almost 90% of the genera. The two main
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s recognized by Groppo et al. in 2012 were upheld, but Morton and Telmer's Rutoideae was
paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
and their Amyridoideae was
polyphyletic
A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
and did not include the type genus. Applehans et al. divided the family into six subfamilies, shown below in the
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 ...
produced in their study. The large subfamily
Zanthoxyloideae
''Zanthoxyloideae'' is a subfamily of the family Rutaceae. Its most notable member is probably Sichuan pepper.
Genera
The division of the subfamily into genera varied, . Genera accepted in a 2021 classification of Rutaceae into subfamilies were: ...
was shown to contain distinct clades, but the authors considered that a revised classification at the tribal level was not yet feasible at the time their paper was published.
Notable species
The family is of great economic importance in warm temperate and subtropical climates for its numerous edible fruits of the genus ''
Citrus
''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
lemon
The lemon (''Citrus'' × ''limon'') is a species of small evergreen tree in the ''Citrus'' genus of the flowering plant family Rutaceae. A true lemon is a hybrid of the citron and the bitter orange. Its origins are uncertain, but some ...
Indian subcontinent
The Indian subcontinent is a physiographic region of Asia below the Himalayas which projects into the Indian Ocean between the Bay of Bengal to the east and the Arabian Sea to the west. It is now divided between Bangladesh, India, and Pakista ...
and elsewhere, as its leaves are used as a spice to flavour dishes. Spices are also made from a number of species in the genus ''
Zanthoxylum
''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the Family (biology), family Rutaceae that are native to warm temperate and Subtropics, subtropical areas worldwide. It is the type gen ...
'', notably
Sichuan pepper
Sichuan pepper (, also known as Sichuanese pepper, Szechuan pepper, Chinese prickly ash, Chinese pepper, Mountain pepper, and ''mala'' pepper, is a spice commonly used in Sichuan cuisine in China, Bhutan and in northeast India. It is called mej ...
.
Other plants are grown in
horticulture
Horticulture (from ) is the art and science of growing fruits, vegetables, flowers, trees, shrubs and ornamental plants. Horticulture is commonly associated with the more professional and technical aspects of plant cultivation on a smaller and mo ...
: ''
Murraya
''Murraya'' () is a genus of flowering plants in the citrus family, Rutaceae. It is distributed in Asia, Australia, and the Pacific Islands.Skimmia'' species, for example. '' Ruta'', ''
Zanthoxylum
''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the Family (biology), family Rutaceae that are native to warm temperate and Subtropics, subtropical areas worldwide. It is the type gen ...
'' and '' Casimiroa'' species are medicinals. Several plants are also used by the
perfume
Perfume (, ) is a mixture of fragrance, fragrant essential oils or aroma compounds (fragrances), Fixative (perfumery), fixatives and solvents, usually in liquid form, used to give the human body, animals, food, objects, and living-spaces an agre ...
industry, such as the Western Australian '' Boronia megastigma''.
The genus '' Pilocarpus'' has species (''P. jaborandi'', and ''P. microphyllus'' from Brazil, and ''P. pennatifolius'' from Paraguay) from which the medicine pilocarpine, used to treat glaucoma, is extracted.