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''Ptelea trifoliata'', commonly known as common hoptree, wafer ash, stinking ash, and skunk bush, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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 ...
in the citrus family (
Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...
). It is
native Native may refer to: People * '' Jus sanguinis'', nationality by blood * '' Jus soli'', nationality by location of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Nat ...
to North America, where it is found in Canada, Mexico, and the United States. It is a
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
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 ...
or
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 ...
, with alternate, trifoliate leaves.


Description

''Ptelea trifoliata'' is a small tree, or often a shrub of a few spreading stems, growing to around tall with a broad crown. The
bark Bark may refer to: Common meanings * 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) Arts and entertainment * ''Bark'' (Jefferson Airplane album), ...
is reddish brown to gray brown, with short horizontal
lenticel A lenticel is a porous tissue consisting of cells with large intercellular spaces in the periderm of the secondarily thickened organs and the Bark (botany), bark of woody stems and roots of gymnosperms and dicotyledonous flowering plants. It func ...
s (warty corky ridges), becoming slightly scaly, The plant has an unpleasant odor and bitter taste. Branchlets are dark reddish brown, shining, covered with small excrescences. The twigs are slender to moderately stout, brown with deep U-shaped leaf scars, and with short, light brown, fuzzy buds. It has thick fleshy roots.


Leaves

Its leaves are alternate and compound with three leaflets, dotted with oil glands. The leaflets are sessile, ovate or oblong, long by broad, pointed at the base, entire or serrate, and gradually pointed at the apex. They are feather-veined, with a prominent midrib and primary veins. They come out of the bud conduplicate and very downy. When fully grown the leaves are dark green and shiny above and paler green beneath. In autumn they turn a rusty yellow. The petioles are stout, long, with an enlarged base.
Stipules 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). They are primarily found among dicots and rare among monocots. Stipules are considered part of the anatomy ...
are absent. The western and southwestern forms have smaller leaves, , than the eastern forms , an adaptation to the drier
climate Climate is the long-term weather pattern in a region, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteoro ...
s in the west.


Flowers

The
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 are small, across, with 4–5 narrow, greenish white petals. The
pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branch ...
are downy. The 4- or 5-part calyx is downy and
imbricate Aestivation or estivation is the positional arrangement of the parts of a flower within a flower bud before it has opened. Aestivation is also sometimes referred to as praefoliation or prefoliation, but these terms may also mean vernation: the ar ...
in bud. The corolla has four or five petals which are white, downy, spreading, hypogynous, and imbricate in bud. The five
stamens 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 filamen ...
alternate with the petals. The
pistillate 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) ...
flowers bear rudimentary
anthers 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 filamen ...
. The filaments are awl-shaped and more-or-less hairy. The anthers are ovate or cordate, two-celled, with cells opening longitudinally. The
ovaries 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 endocr ...
are superior, hairy, abortive in the staminate flowers, two to three-celled. The style is short, the stigma 2- or 3-lobed, with two ovules per cell. Fertile and sterile flowers are produced together in terminal, spreading, compound
cymes In botany, an inflorescence is a group or cluster of flowers arranged on a plant's stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a main axis ( ped ...
—the sterile being usually fewer and falling after the anther cells mature. Flowers are produced in May and June. Some find the odor unpleasant but to others the plant has a delicious scent.


Fruit

The
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
is a round wafer-like papery
samara Samara, formerly known as Kuybyshev (1935–1991), is the largest city and administrative centre of Samara Oblast in Russia. The city is located at the confluence of the Volga and the Samara (Volga), Samara rivers, with a population of over 1.14 ...
, across, light brown, and two-seeded. The fruit ripens in October, and is held on the tree until high winds shake them loose in the early winter.


Wood

Its wood is yellow brown; heavy, hard, close-grained, satiny. The
specific gravity Relative density, also called specific gravity, is a dimensionless quantity defined as the ratio of the density (mass of a unit volume) of a substance to the density of a given reference material. Specific gravity for solids and liquids is nea ...
is 0.8319; weight per cubic foot is .


Gallery

File:Ptelea trifoliata Kentucky.jpg, Flowers of ''Ptelea trifoliata'' File:Ptelea trifoliata MN 2007.JPG, Multi-trunk tree form


Taxonomy

While ''Ptelea trifoliata'' is most often treated as a single species with
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
and/or
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
in different distribution ranges, some botanists treat the various hoptrees as a group of four or more closely related species: *''P. trifoliata'' subsp. ''trifoliata'' – common hoptree or eastern hoptree; eastern Canada & United States (U.S.), central U.S. **''P. trifoliata'' subsp. ''trifoliata'' var. ''trifoliata'' – eastern Canada & (U.S.), central U.S. **''P. trifoliata'' subsp ''trifoliata'' var. ''mollis'' Torr. & A. Gray – eastern and central U.S. *''P. trifoliata'' subsp. ''angustifolia'' (Benth.) V.L.Bailey – south-central U.S. **''P. trifoliata'' subsp. ''angustifolia'' var. ''angustifolia'' (Benth.) M.E.Jones (''P. angustifolia'', ''P. lutescens'') – narrowleaf hoptree; south-central U.S. **''P. trifoliata'' subsp. ''angustifolia'' var. ''persicifolia'' (Greene) V.L.Bailey – south-central U.S. *''P. trifoliata'' subsp. ''pallida'' (Greene) V.L.Bailey – pallid hoptree, south-central and southwest U.S. **''P. trifoliata'' subsp. ''pallida'' var. ''pallida'' (Greene) V.L.Bailey – southwest U.S. **''P. trifoliata'' subsp. ''pallida'' var. ''cognata'' (Greene) Kearney & Peebles – southwest U.S. **''P. trifoliata'' subsp. ''pallida'' var. ''confinis'' (Greene) V.L.Bailey – south-central and southwest U.S. **''P. trifoliata'' subsp. ''pallida'' var. ''lutescens'' – southwest U.S. *''P. trifoliata'' subsp. ''polyadenia'' (Greene) V.L.Bailey – pallid hoptree, south-central and southwest U.S. *''P. trifoliata'' var. ''baldwinii'' (Torr. & A.Gray) D.B.Ward (''P. baldwinii'') The specific epithet "''trifoliata''" refers to the three-parted compound
leaf A leaf (: leaves) is a principal appendage of the plant stem, 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 leav ...
. Other common names for this shrub include stinking prairie bush, Carolina shrub-trefoil, tree-trefoil, swamp dogwood, ague bark, paleleaf hoptree, prairie-grub, prickaway-anise, quinine tree, sang-tree, water-ash, western hoptree, wingseed, and woolly hoptree.


Distribution and habitat

''Ptelea trifoliata'' is native to North America, where its northern limits are in
Ontario Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
and
Quebec Quebec is Canada's List of Canadian provinces and territories by area, largest province by area. Located in Central Canada, the province shares borders with the provinces of Ontario to the west, Newfoundland and Labrador to the northeast, ...
, Canada. It is native through much of the eastern and southwestern
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
, although it is absent from some areas of the
Upper Midwest The Upper Midwest is a northern subregion of the U.S. Census Bureau's Midwestern United States. Although the exact boundaries are not uniformly agreed upon, the region is usually defined to include the states of Iowa, Michigan, Minnesota and Wi ...
and is rare in much of
New England New England is a region consisting of six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York (state), New York to the west and by the ...
. Its southern limits are in
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 ...
. It has a wide-ranging natural habitat. In the
Southeastern United States The Southeastern United States, also known as the American Southeast or simply the Southeast, is a geographical List of regions in the United States, region of the United States located in the eastern portion of the Southern United States and t ...
it is most often found in rocky forests, in both moist and dry soil, often associated with calcareous or mafic substrates. In the
Midwest The Midwestern United States (also referred to as the Midwest, the Heartland or the American Midwest) is one of the four census regions defined by the United States Census Bureau. It occupies the northern central part of the United States. It ...
, habitats include forests, savannas, prairies, glades, and sand dunes. In
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
it is common in canyons.


Ecology

Larva of the giant swallowtail butterfly '' Papilio cresphontes'' feed on the leaves.
Treehopper Treehoppers (more precisely typical treehoppers to distinguish them from the Aetalionidae) and thorn bugs are members of the family Membracidae, a group of insects related to the cicadas and the leafhoppers. About 3,200 species of treehoppers i ...
s of the genus '' Enchenopa'' infest the branches, laying white-frothy masses of eggs on the branch undersides. Several ant species tend to the treehoppers, including '' Camponotus pennsylvanicus'', '' Formica montana'', and '' Formica subsericea''. Several bee species have been documented visiting the flowers of wafer ash, including '' Agapostemon virescens'', '' Andrena commoda'', '' Andrena crataegi'', '' Andrena cressonii'', ''
Apis mellifera The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for 'bee', and ''mellifera'' is the Latin for 'honey-bearing' or 'honey-carrying', ...
'', '' Bombus auricomus'', '' Bombus bimaculatus'', '' Bombus impatiens'', '' Ceratina calcarata'', '' Ceratina dupla'', '' Ceratina mikmaqi'', and '' Lasioglossum imitatum''.


Uses

It has several Native American uses as a seasoning and as an
herbal medicine Herbal medicine (also called herbalism, phytomedicine or phytotherapy) is the study of pharmacognosy and the use of medicinal plants, which are a basis of traditional medicine. Scientific evidence for the effectiveness of many herbal treatments ...
for different ailments. Numerous
cultivars A cultivar is a kind of cultivated plant that people have selected for desired traits and which retains those traits when propagated. Methods used to propagate cultivars include division, root and stem cuttings, offsets, grafting, tissue cult ...
have been developed for ornamental use in parks and gardens. The cultivar 'Aurea' with golden leaves has gained the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr ...
's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. It includes the full range of cultivated p ...
. German immigrants to Texas in the 19th century used its seeds in place of
hops Hops are the flowers (also called seed cones or strobiles) of the hop plant ''Humulus lupulus'', a member of the Cannabaceae family of flowering plants. They are used primarily as a bittering, flavouring, and stability agent in beer, to whic ...
in the beer-making process, lending the species its common name.


References

{{Authority control Zanthoxyloideae Trees of Northern America Flora of California Flora of Central Mexico Plants described in 1753 Taxa named by Carl Linnaeus Medicinal plants Garden plants of North America