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Elm
Elm
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Elm
Temporal range: Maastrichtian[1]–Recent
U. minor, East Coker
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Rosales
Family: Ulmaceae
Genus: Ulmus
L.
Species

See

Elms are deciduous and semi-deciduous trees comprising the genus Ulmus in the family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical-montane regions of North America and Eurasia, ranging southward in Western Asia to Iran, in Africa to Libya, and in Southeast Asia into Indonesia.[2]

Elms are components of many kinds of natural forests. Moreover, during the 19th and early 20th centuries, many species and cultivars were also planted as ornamental street, garden, and park trees in Europe, North America, and parts of the Southern Hemisphere, notably Australasia. Some individual elms reached great size and age. However, in recent decades, most mature elms of European or North American origin have died from Dutch elm disease, caused by a microfungus dispersed by bark beetles. In response, disease-resistant cultivars have been developed, capable of restoring the elm to forestry and landscaping.

Description

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The genus is hermaphroditic, having apetalous perfect flowers which are wind-pollinated. Elm leaves are alternate, with simple, single- or, most commonly, doubly serrate margins, usually asymmetric at the base and acuminate at the apex. The fruit is a round wind-dispersed samara flushed with chlorophyll, facilitating photosynthesis before the leaves emerge.[3] The samarae are very light, those of British elms numbering around 50,000 to the pound (454 g).[4] (Very rarely anomalous samarae occur with more than two wings.[5]) All species are tolerant of a wide range of soils and pH levels but, with few exceptions, demand good drainage. The elm tree can grow to great height, the American elm in excess of 30 metres (98 feet),[6] often with a forked trunk creating a vase profile.

Taxonomy

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There are about 30 to 40 species of Ulmus; the ambiguity in number results from difficulty in delineating species, owing to the ease of hybridization between them and the development of local seed-sterile vegetatively propagated microspecies in some areas, mainly in the field elm (Ulmus minor) group. Oliver Rackham[7] describes Ulmus as the most critical genus in the entire British flora, adding that 'species and varieties are a distinction in the human mind rather than a measured degree of genetic variation'. Eight species are endemic to North America and three to Europe, but the greatest diversity is in Asia with approximately two dozen species.[8] The oldest fossils of Ulmus are leaves dating Paleocene, found across the Northern Hemisphere.[9]

The classification adopted in the List of elm species is largely based on that established by Brummitt.[10] A large number of synonyms have accumulated over the last three centuries; their currently accepted names can be found in the list of Elm synonyms and accepted names.

Botanists who study elms and argue over elm identification and classification are called "pteleologists", from the Greek πτελέα (elm).[11]

As part of the order Urticales, they are distantly related to cannabis, mulberries, figs, hops, and nettles.

Ecology

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Propagation

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A rooted cutting of European white elm (July)

Elm propagation methods vary according to elm type and location, and the plantsman's needs. Native species may be propagated by seed. In their natural setting, native species, such as wych elm and European white elm in central and northern Europe and field elm in southern Europe, set viable seed in "favourable" seasons. Optimal conditions occur after a late warm spring.[12] After pollination, seeds of spring-flowering elms ripen and fall at the start of summer (June); they remain viable for only a few days. They are planted in sandy potting soil at a depth of 1 cm, and germinate in three weeks. Slow-germinating American elm will remain dormant until the second season.[13] Seeds from autumn-flowering elms ripen in the fall and germinate in the spring.[13] Since elms may hybridize within and between species, seed propagation entails a hybridisation risk. In unfavourable seasons, elm seeds are usually sterile. Elms outside their natural range, such as English elm U. minor 'Atinia', and elms unable to pollinate because pollen sources are genetically identical, are sterile and are propagated by vegetative reproduction. Vegetative reproduction is also used to produce genetically identical elms (clones). Methods include the winter transplanting of root suckers; taking hardwood cuttings from vigorous one-year-old shoots in late winter,[14] taking root cuttings in early spring; taking softwood cuttings in early summer;[15] grafting; ground and air layering; and micropropagation. A bottom heat of 18 °C[16] and humid conditions are maintained for hard- and softwood cuttings. The transplanting of root suckers remains the easiest and most common propagation method for European field elm and its hybrids. For specimen urban elms, grafting to wych-elm rootstock may be used to eliminate suckering or to ensure stronger root growth. The mutant-elm cultivars are usually grafted, the "weeping" elms 'Camperdown' and 'Horizontalis' at 2–3 m (7–10 ft), the dwarf cultivars 'Nana' and 'Jacqueline Hillier' at ground level. Since the Siberian elm is drought tolerant, in dry countries, new varieties of elm are often root-grafted onto this species.[17]

Associated organisms

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Pests and diseases

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Dutch elm disease

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Golden elm tree with Dutch elm disease

Dutch elm disease (DED) devastated elms throughout Europe and much of North America in the second half of the 20th century. It derives its name "Dutch" from the first description of the disease and its cause in the 1920s by Dutch botanists Bea Schwarz and Christina Johanna Buisman. Owing to its geographical isolation and effective quarantine enforcement, Australia has so far remained unaffected by DED, as have the provinces of Alberta and British Columbia in western Canada.

DED is caused by a microfungus transmitted by two species of Scolytus elm-bark beetles, which act as vectors. The disease affects all species of elms native to North America and Europe, but many Asiatic species have evolved antifungal genes and are resistant. Fungal spores, introduced into wounds in the tree caused by the beetles, invade the xylem or vascular system. The tree responds by producing tyloses, effectively blocking the flow from roots to leaves. Woodland trees in North America are not quite as susceptible to the disease because they usually lack the root grafting of the urban elms and are somewhat more isolated from each other. In France, inoculation with the fungus of over 300 clones of the European species failed to find a single variety that possessed of any significant resistance.

The first, less aggressive strain of the disease fungus, Ophiostoma ulmi, arrived in Europe from Asia in 1910, and was accidentally introduced to North America in 1928. It was steadily weakened by viruses in Europe and had all but disappeared by the 1940s. However, the disease had a much greater and longer-lasting impact in North America, owing to the greater susceptibility of the American elm, Ulmus americana, which masked the emergence of the second, far more virulent strain of the disease Ophiostoma novo-ulmi. It appeared in the United States sometime in the 1940s, and was originally believed to be a mutation of O. ulmi. Limited gene flow from O. ulmi to O. novo-ulmi was probably responsible for the creation of the North American subspecies O. novo-ulmi subsp. americana. It was first recognized in Britain in the early 1970s, believed to have been introduced via a cargo of Canadian rock elm destined for the boatbuilding industry, and rapidly eradicated most of the mature elms from western Europe. A second subspecies, O. novo-ulmi subsp. novo-ulmi, caused similar devastation in Eastern and Central Europe. This subspecies, which was introduced to North America, and like O. ulmi, is thought to have originated in Asia. The two subspecies have now hybridized in Europe where their ranges have overlapped.[18] The hypothesis that O. novo-ulmi arose from a hybrid of the original O. ulmi and another strain endemic to the Himalayas, Ophiostoma himal-ulmi, is now discredited.[19]

No sign indicates the current pandemic is waning, and no evidence has been found of a susceptibility of the fungus to a disease of its own caused by d-factors: naturally occurring virus-like agents that severely debilitated the original O. ulmi and reduced its sporulation.[20]

Elm phloem necrosis

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Elm phloem necrosis (elm yellows) is a disease of elm trees that is spread by leafhoppers or by root grafts.[21] This very aggressive disease, with no known cure, occurs in the Eastern United States, southern Ontario in Canada, and Europe. It is caused by phytoplasmas that infect the phloem (inner bark) of the tree.[22] Infection and death of the phloem effectively girdles the tree and stops the flow of water and nutrients. The disease affects both wild-growing and cultivated trees. Occasionally, cutting the infected tree before the disease completely establishes itself and cleanup and prompt disposal of infected matter has resulted in the plant's survival via stump sprouts.

Insects

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The elm leaf beetle

Most serious of the elm pests is the elm leaf beetle Xanthogaleruca luteola, which can decimate foliage, although rarely with fatal results. The beetle was accidentally introduced to North America from Europe. Another unwelcome immigrant to North America is the Japanese beetle Popillia japonica. In both instances, the beetles cause far more damage in North America owing to the absence of the predators in their native lands. In Australia, introduced elm trees are sometimes used as food plants by the larvae of hepialid moths of the genus Aenetus. These burrow horizontally into the trunk then vertically down.[23][24] Circa 2000, the Asian Zig-zag sawfly Aproceros leucopoda appeared in Europe and North America, although in England, its impact has been minimal and it is no longer monitored.[25]

Birds

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Sapsucker woodpeckers have a great love of young elm trees.[26]

The ball-headed graft narvan elm, Ulmus minor 'Umbraculifera', cultivated in Persia and widely planted in central Asia
Lafayette Street in Salem, Massachusetts - an example of the high-tunneled effects of Ulmus americana avenues once common in New England (colorized postcard, 1910)

Cultivation

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Camperdown elm (Ulmus glabra 'Camperdownii'), cultivated in Prospect Park, Brooklyn, New York
An avenue of elm trees in Fitzroy Gardens, Melbourne

One of the earliest of ornamental elms was the ball-headed graft narvan elm, Ulmus minor 'Umbraculifera', cultivated from time immemorial in Persia as a shade tree and widely planted in cities through much of south-west and central Asia. From the 18th century to the early 20th century, elms, whether species, hybrids, or cultivars, were among the most widely planted ornamental trees in both Europe and North America. They were particularly popular as a street tree in avenue plantings in towns and cities, creating high-tunnelled effects. Their quick growth and variety of foliage and forms,[27] their tolerance of air-pollution, and the comparatively rapid decomposition of their leaf litter in the fall were further advantages.

In North America, the species most commonly planted was the American elm (U. americana), which had unique properties that made it ideal for such use - rapid growth, adaptation to a broad range of climates and soils, strong wood, resistance to wind damage, and vase-like growth habit requiring minimal pruning. In Europe, the wych elm (U. glabra) and the field elm (U. minor) were the most widely planted in the countryside, the former in northern areas including Scandinavia and northern Britain, the latter further south. The hybrid between these two, Dutch elm (U. × hollandica), occurs naturally and was also commonly planted. In much of England, the English elm later came to dominate the horticultural landscape. Most commonly planted in hedgerows, it sometimes occurred in densities over 1000/km2. In south-eastern Australia and New Zealand, large numbers of English and Dutch elms, as well as other species and cultivars, were planted as ornamentals following their introduction in the 19th century, while in northern Japan Japanese elm (U. davidiana var. japonica) was widely planted as a street tree. From about 1850 to 1920, the most prized small ornamental elm in parks and gardens was the 'Camperdown' elm (U. glabra 'Camperdownii'), a contorted, weeping cultivar of the wych elm grafted on to a nonweeping elm trunk to give a wide, spreading, and weeping fountain shape in large garden spaces.

In northern Europe, elms were among the few trees tolerant of saline deposits from sea spray, which can cause "salt-burning" and die-back. This tolerance made elms reliable both as shelterbelt trees exposed to sea wind, in particular along the coastlines of southern and western Britain[28][29] and in the Low Countries, and as trees for coastal towns and cities.[30]

This belle époque lasted until the First World War, when the elm began its slide into horticultural decline. The impact of the hostilities on Germany, the origin of at least 40 cultivars, coincided with an outbreak of the early strain of DED, Ophiostoma ulmi. The devastation caused by the Second World War, and the demise in 1944 of the huge Späth nursery in Berlin, only accelerated the process. The outbreak of the new, three times more virulent, strain of DED Ophiostoma novo-ulmi in the late 1960s, brought the tree to its nadir.

Since around 1990, the elm has enjoyed a renaissance through the successful development in North America and Europe of cultivars highly resistant to DED.[3] Consequently, the total number of named cultivars, ancient and modern, now exceeds 300, although many of the older clones, possibly over 120, have been lost to cultivation. Some of the latter, however, were by today's standards inadequately described or illustrated before the pandemic, and a number may survive, or have regenerated, unrecognised. Enthusiasm for the newer clones often remains low owing to the poor performance of earlier, supposedly disease-resistant Dutch trees released in the 1960s and 1970s. In the Netherlands, sales of elm cultivars slumped from over 56,000 in 1989 to just 6,800 in 2004,[31] whilst in the UK, only four of the new American and European releases were commercially available in 2008.

A row of Princeton elm trees, which are moderately resistant to Dutch elm disease, at Scripps College in Claremont, California.[32]

Efforts to develop DED-resistant cultivars began in the Netherlands in 1928 and continued, uninterrupted by World War II, until 1992.[33] Similar programmes were initiated in North America (1937), Italy (1978) and Spain (1986). Research has followed two paths:

Species and species cultivars

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In North America, careful selection has produced a number of trees resistant not only to DED, but also to the droughts and cold winters that occur on the continent. Research in the United States has concentrated on the American elm (U. americana), resulting in the release of DED-resistant clones, notably the cultivars 'Valley Forge' and 'Jefferson'. Much work has also been done into the selection of disease-resistant Asiatic species and cultivars.[34][35]

In 1993, Mariam B. Sticklen and Mark G. Bolyard reported the results of experiments funded by the US National Park Service and conducted at Michigan State University in East Lansing that were designed to apply genetic engineering techniques to the development of DED-resistant strains of American elm trees.[36] In 2007, A. E. Newhouse and F. Schrodt of the State University of New York College of Environmental Science and Forestry in Syracuse reported that young transgenic American elm trees had shown reduced DED symptoms and normal mycorrhizal colonization.[37]

In Europe, the European white elm (U. laevis) has received much attention. While this elm has little innate resistance to DED, it is not favoured by the vector bark beetles. Thus it becomes colonized and infected only when no other elms are available, a rare situation in western Europe. Research in Spain has suggested that it may be the presence of a triterpene, alnulin, which makes the tree bark unattractive to the beetle species that spread the disease.[38] This possibility, though, has not been conclusively proven.[39] More recently, field elms Ulmus minor highly resistant to DED have been discovered in Spain, and form the basis of a major breeding programme.[40]

Hybrid cultivars

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Owing to their innate resistance to DED, Asiatic species have been crossed with European species, or with other Asiatic elms, to produce trees that are both highly resistant to disease and tolerant of native climates. After a number of false dawns in the 1970s, this approach has produced a range of reliable hybrid cultivars now commercially available in North America and Europe.[41][42][43][44][45][46][47] Disease resistance is invariably carried by the female parent.[48]

Some of these cultivars, notably those with the Siberian elm (Ulmus pumila) in their ancestry, lack the forms for which the iconic American elm and English elm were prized. Moreover, several exported to northwestern Europe have proven unsuited to the maritime climate conditions there, notably because of their intolerance of anoxic conditions resulting from ponding on poorly drained soils in winter. Dutch hybridizations invariably included the Himalayan elm (Ulmus wallichiana) as a source of antifungal genes and have proven more tolerant of wet ground; they should also ultimately reach a greater size. However, the susceptibility of the cultivar 'Lobel', used as a control in Italian trials, to elm yellows has now (2014) raised a question mark over all the Dutch clones.[49]

Several highly resistant Ulmus cultivars have been released since 2000 by the Institute of Plant Protection in Florence, most commonly featuring crosses of the Dutch cultivar 'Plantijn' with the Siberian elm to produce resistant trees better adapted to the Mediterranean climate.[42]

Cautions regarding novel cultivars

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Elms take many decades to grow to maturity, and as the introduction of these disease-resistant cultivars is relatively recent, their long-term performance and ultimate size and form cannot be predicted with certainty. The National Elm Trial in North America, begun in 2005, is a nationwide trial to assess strengths and weaknesses of the 19 leading cultivars raised in the US over a 10-year period; European cultivars have been excluded.[50] Meanwhile, in Europe, American and European cultivars are being assessed in field trials started in 2000 by the UK charity Butterfly Conservation.[51]

Landscaped parks

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Central Park

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American elm trees along the Mall and Literary Walk in New York City's Central Park (2013)

The oldest American elm trees in New York City's Central Park were planted in the 1860s by Frederick Law Olmsted, making them among the oldest stands of American elms in the world. Along the Mall and Literary Walk four lines of American elms stretch over the walkway forming a cathedral-like covering. A part of New York City's urban ecology, the elms improve air and water quality, reduce erosion and flooding, and decrease air temperatures during warm days.[52]

While the stand is still vulnerable to DED, in the 1980s the Central Park Conservancy undertook aggressive countermeasures such as heavy pruning and removal of extensively diseased trees. These efforts have largely been successful in saving the majority of the trees, although several are still lost each year. Younger American elms that have been planted in Central Park since the outbreak are of the DED-resistant 'Princeton' and 'Valley Forge' cultivars.[53]

National Mall

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Rows of American elm trees line a path south of the Lincoln Memorial Reflecting Pool on the National Mall in Washington, DC (November 11, 2006)

Several rows of American elm trees that the National Park Service (NPS) first planted during the 1930s line much of the 1.9-mile-length (3.1-kilometer) of the National Mall in Washington, DC. DED first appeared on the trees during the 1950s and reached a peak in the 1970s. The NPS used a number of methods to control the epidemic, including sanitation, pruning, injecting trees with fungicide, and replanting with DED-resistant cultivars. The NPS combated the disease's local insect vector, the smaller European elm bark beetle (Scolytus multistriatus), by trapping and by spraying with insecticides. As a result, the population of American elms planted on the Mall and its surrounding areas has remained intact for more than 80 years.[54]

Uses

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Wood

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Elm wood
Elm in boatbuilding: John Constable, Boat-Building Near Flatford Mill, 1815 (landscape with hybrid elms Ulmus × hollandica[12])
English longbow of elm

Elm wood is valued for its interlocking grain, and consequent resistance to splitting, with significant uses in wagon-wheel hubs, chair seats, and coffins. The bodies of Japanese Taiko drums are often cut from the wood of old elm trees, as the wood's resistance to splitting is highly desired for nailing the skins to them, and a set of three or more is often cut from the same tree. The elm's wood bends well and distorts easily. The often long, straight trunks were favoured as a source of timber for keels in ship construction. Elm is also prized by bowyers; of the ancient bows found in Europe, a large portion are elm. During the Middle Ages, elm was also used to make longbows if yew was unavailable.

The first written references to elm occur in the Linear B lists of military equipment at Knossos in the Mycenaean period. Several of the chariots are of elm ("πτε-ρε-ϝα", pte-re-wa), and the lists twice mention wheels of elmwood.[55] Hesiod says that ploughs in Ancient Greece were also made partly of elm.[56]

The density of elm wood varies between species, but averages around 560 kg/m3.[57]

Elm wood is also resistant to decay when permanently wet, and hollowed trunks were widely used as water pipes during the medieval period in Europe. Elm was also used as piers in the construction of the original London Bridge, but this resistance to decay in water does not extend to ground contact.[57]

Viticulture

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The Romans, and more recently, Italians, planted elms in vineyards as supports for vines. Lopped at 3 m, the elms' quick growth, twiggy lateral branches, light shade, and root suckering made them ideal trees for this purpose. The lopped branches were used for fodder and firewood.[58] Ovid in his Amores characterizes the elm as "loving the vine": ulmus amat vitem, vitis non deserit ulmum (the elm loves the vine, the vine does not desert the elm),[59] and the ancients spoke of the "marriage" between elm and vine.[60]

Medicinal products

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The mucilaginous inner bark of the slippery elm (Ulmus rubra) has long been used as a demulcent, and is still produced commercially for this purpose in the US with approval for sale as a nutritional supplement by the Food and Drug Administration.[61]

Fodder

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Elms also have a long history of cultivation for fodder, with the leafy branches cut to feed livestock. The practice continues today in the Himalaya, where it contributes to serious deforestation.[62]

Biomass

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As fossil fuel resources diminish, increasing attention is being paid to trees as sources of energy. In Italy, the Istituto per la Protezione delle Piante is (2012) in the process of releasing to commerce very fast-growing elm cultivars, able to increase in height by more than 2 metres (6.6 feet) per year.[63]

Food

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Elm bark, cut into strips and boiled, sustained much of the rural population of Norway during the great famine of 1812. The seeds are particularly nutritious, containing 45% crude protein, and less than 7% fibre by dry mass.[64]

Internal mill-wheel of elm, De Hoop mill, Oldebroek, Netherlands

Alternative medicine

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Elm has been listed as one of the 38 substances that are used to prepare Bach flower remedies,[65] a kind of alternative medicine.

Bonsai

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Chinese Elm Ulmus parvifolia bonsai

Chinese elm (Ulmus parvifolia) is a popular choice for bonsai owing to its tolerance of severe pruning.

Genetic resource conservation

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In 1997, a European Union elm project was initiated, its aim to coordinate the conservation of all the elm genetic resources of the member states and, among other things, to assess their resistance to Dutch elm disease. Accordingly, over 300 clones were selected and propagated for testing.[66][67][68]

Culture

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Notable elm trees

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Many elm trees of various kinds have attained great size or otherwise become particularly noteworthy.

In art

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Many artists have admired elms for the ease and grace of their branching and foliage, and have painted them with sensitivity. Elms are a recurring element in the landscapes and studies of, for example, John Constable, Ferdinand Georg Waldmüller, Alfred East, George Clausen, Frederick Childe Hassam, Karel Klinkenberg,[69] and George Inness.

In mythology and literature

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Achilles and Scamander
Dryad

In Greek mythology, the nymph Ptelea (Πτελέα, Elm) was one of the eight hamadryads, nymphs of the forest and daughters of Oxylos and Hamadryas.[72] In his Hymn to Artemis, poet Callimachus (third century BC) tells how, at the age of three, the infant goddess Artemis practised her newly acquired silver bow and arrows, made for her by Hephaestus and the Cyclopes, by shooting first at an elm, then at an oak, before turning her aim on a wild animal:

πρῶτον ἐπὶ πτελέην, τὸ δὲ δεύτερον ἧκας ἐπὶ δρῦν, τὸ τρίτον αὖτ᾽ ἐπὶ θῆρα.[73]

The first reference in literature to elms occurs in the Iliad. When Eetion, father of Andromache, is killed by Achilles during the Trojan War, the mountain nymphs plant elms on his tomb ("περί δὲ πτελέας ἐφύτευσαν νύμφαι ὀρεστιάδες, κoῦραι Διὸς αἰγιόχoιo").[74] Also in the Iliad, when the River Scamander, indignant at the sight of so many corpses in his water, overflows and threatens to drown Achilles, the latter grasps a branch of a great elm in an attempt to save himself ("ὁ δὲ πτελέην ἕλε χερσὶν εὐφυέα μεγάλην").[75]

The nymphs also planted elms on the tomb in the Thracian Chersonese of "great-hearted Protesilaus" ("μεγάθυμου Πρωτεσιλάου"), the first Greek to fall in the Trojan War. These elms grew to be the tallest in the known world, but when their topmost branches saw far off the ruins of Troy, they immediately withered, so great still was the bitterness of the hero buried below, who had been loved by Laodamia and slain by Hector.[76][77][78] The story is the subject of a poem by Antiphilus of Byzantium (first century AD) in the Palatine Anthology:

Θεσσαλὲ Πρωτεσίλαε, σὲ μὲν πολὺς ᾄσεται αἰών,
Tρoίᾳ ὀφειλoμένoυ πτώματος ἀρξάμενoν•
σᾶμα δὲ τοι πτελέῃσι συνηρεφὲς ἀμφικoμεῦση
Nύμφαι, ἀπεχθoμένης Ἰλίoυ ἀντιπέρας.
Δένδρα δὲ δυσμήνιτα, καὶ ἤν ποτε τεῖχoς ἴδωσι
Tρώϊον, αὐαλέην φυλλοχoεῦντι κόμην.
ὅσσoς ἐν ἡρώεσσι τότ᾽ ἦν χόλoς, oὗ μέρoς ἀκμὴν
ἐχθρὸν ἐν ἀψύχoις σώζεται ἀκρέμoσιν.[79]
[:Thessalian Protesilaos, a long age shall sing your praises,
Of the destined dead at Troy the first;
Your tomb with thick-foliaged elms they covered,
The nymphs, across the water from hated Ilion.
Trees full of anger; and whenever that wall they see,
Of Troy, the leaves in their upper crown wither and fall.
So great in the heroes was the bitterness then, some of which still
Remembers, hostile, in the soulless upper branches.]

Protesilaus had been king of Pteleos (Πτελεός) in Thessaly, which took its name from the abundant elms (πτελέoι) in the region.[80]

Elms occur often in pastoral poetry, where they symbolise the idyllic life, their shade being mentioned as a place of special coolness and peace. In the first Idyll of Theocritus (third century BC), for example, the goatherd invites the shepherd to sit "here beneath the elm" ("δεῦρ' ὑπὸ τὰν πτελέαν") and sing. Beside elms, Theocritus places "the sacred water" ("το ἱερὸν ὕδωρ") of the Springs of the Nymphs and the shrines to the nymphs.[81]

The Sibyl and Aeneas

Aside from references literal and metaphorical to the elm and vine theme, the tree occurs in Latin literature in the Elm of Dreams in the Aeneid.[82] When the Sibyl of Cumae leads Aeneas down to the Underworld, one of the sights is the Stygian Elm:

In medio ramos annosaque bracchia pandit
ulmus opaca, ingens, quam sedem somnia vulgo
uana tenere ferunt, foliisque sub omnibus haerent.
[:Spreads in the midst her boughs and agéd arms
an elm, huge, shadowy, where vain dreams, 'tis said,
are wont to roost them, under every leaf close-clinging.]

Virgil refers to a Roman superstition (vulgo) that elms were trees of ill-omen because their fruit seemed to be of no value.[83] It has been noted[84] that two elm-motifs have arisen from classical literature: (1) the 'Paradisal Elm' motif, arising from pastoral idylls and the elm-and-vine theme, and (2) the 'Elm and Death' motif, perhaps arising from Homer's commemorative elms and Virgil's Stygian Elm. Many references to elm in European literature from the Renaissance onwards fit into one or other of these categories.

There are two examples of pteleogenesis (:birth from elms) in world myths. In Germanic and Scandinavian mythology the first woman, Embla, was fashioned from an elm,[85] while in Japanese mythology Kamuy Fuchi, the chief goddess of the Ainu people, "was born from an elm impregnated by the Possessor of the Heavens".[86]

Under the elm, Brighton, 2006

The elm occurs frequently in English literature, one of the best known instances being in Shakespeare's A Midsummer Night's Dream, where Titania, Queen of the Fairies, addresses her beloved Nick Bottom using an elm-simile. Here, as often in the elm-and-vine motif, the elm is a masculine symbol:

Sleep thou, and I will wind thee in my arms.
... the female Ivy so
Enrings the barky fingers of the Elm.
O, how I love thee! how I dote on thee![87]

Another of the most famous kisses in English literature, that of Paul and Helen at the start of Forster's Howards End, is stolen beneath a great wych elm.

The elm tree is also referenced in children's literature. An Elm Tree and Three Sisters by Norma Sommerdorf is a children's book about three young sisters who plant a small elm tree in their backyard.[88]

In politics

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The cutting of the elm was a diplomatic altercation between the kings of France and England in 1188, during which an elm tree near Gisors in Normandy was felled.[89]

In politics, the elm is associated with revolutions. In England after the Glorious Revolution of 1688, the final victory of parliamentarians over monarchists, and the arrival from Holland, with William III and Mary II, of the Dutch elm hybrid, planting of this cultivar became a fashion among enthusiasts of the new political order.[90][91]

In the American Revolution, the Liberty Tree was an American white elm in Boston, Massachusetts, in front of which, from 1765, the first resistance meetings were held against British attempts to tax the American colonists without democratic representation. When the British, knowing that the tree was a symbol of rebellion, felled it in 1775, the Americans took to widespread Liberty Elm planting, and sewed elm symbols on to their revolutionary flags.[92][93] Elm planting by American Presidents later became something of a tradition.

In the French Revolution, too, Les arbres de la liberté (Liberty Trees), often elms, were planted as symbols of revolutionary hopes, the first in Vienne, Isère, in 1790, by a priest inspired by the Boston elm.[92] L'Orme de La Madeleine (:the Elm of La Madeleine), Faycelles, Département de Lot, planted around 1790 and surviving to this day, was a case in point.[94] By contrast, a famous Parisian elm associated with the Ancien Régime, L'Orme de Saint-Gervais in the Place St-Gervais, was felled by the revolutionaries; church authorities planted a new elm in its place in 1846, and an early 20th-century elm stands on the site today.[95] Premier Lionel Jospin, obliged by tradition to plant a tree in the garden of the Hôtel Matignon, the official residence and workplace of Prime Ministers of France, insisted on planting an elm, so-called 'tree of the Left', choosing the new disease-resistant hybrid 'Clone 762' (Ulmus 'Wanoux' = Vada).[96] In the French Republican Calendar, in use from 1792 to 1806, the 12th day of the month Ventôse (= 2 March) was officially named "jour de l'Orme", Day of the Elm.

Liberty Elms were also planted in other countries in Europe to celebrate their revolutions, an example being L'Olmo di Montepaone, L'Albero della Libertà (:the Elm of Montepaone, Liberty Tree) in Montepaone, Calabria, planted in 1799 to commemorate the founding of the democratic Parthenopean Republic, and surviving until it was brought down by a recent storm (it has since been cloned and 'replanted').[97] After the Greek Revolution of 1821–32, a thousand young elms were brought to Athens from Missolonghi, "Sacred City of the Struggle" against the Turks and scene of Lord Byron's death, and planted in 1839–40 in the National Garden.[98][99] In an ironic development, feral elms have spread and invaded the grounds of the abandoned Greek royal summer palace at Tatoi in Attica.

In a chance event linking elms and revolution, on the morning of his execution (30 January 1649), walking to the scaffold at the Palace of Whitehall, King Charles I turned to his guards and pointed out, with evident emotion, an elm near the entrance to Spring Gardens that had been planted by his brother in happier days. The tree was said to be still standing in the 1860s.[100]

In local history and place names

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The name of what is now the London neighbourhood of Seven Sisters is derived from seven elms which stood there at the time when it was a rural area, planted a circle with a walnut tree at their centre, and traceable on maps back to 1619.[101][102]

See also

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References

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Monographs

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  • Richens, R. H. (1983). Elm. Cambridge University Press. ISBN 0-521-24916-3. A scientific, historical and cultural study, with a thesis on elm-classification, followed by a systematic survey of elms in England, region by region. Illustrated.
  • Heybroek, H. M., Goudzwaard, L, Kaljee, H. (2009). Iep of olm, karakterboom van de Lage Landen (:Elm, a tree with character of the Low Countries). KNNV, Uitgeverij. ISBN 9789050112819. A history of elm planting in the Netherlands, concluding with a 40 – page illustrated review of all the DED – resistant cultivars in commerce in 2009.

Further reading

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Elms comprise the genus Ulmus in the family , consisting of approximately 25-30 species of mostly trees native to temperate and boreal regions of the , with greatest diversity in central and northern . These trees typically exhibit vase-shaped or spreading crowns, serrated leaves, and small wind-pollinated flowers producing winged samaras, adapting them to a range of habitats from floodplains to uplands. Valued historically for their durable wood in furniture, , and , as well as for shade and ornamental planting in urban landscapes, elms played key ecological roles in supporting and stabilizing riparian zones. However, the introduction of in the early 20th century, caused by the fungi Ophiostoma ulmi and O. novo-ulmi and vectored by elm bark beetles, has decimated populations, particularly of susceptible species like the American elm (U. americana), leading to the loss of tens of millions of mature trees across and . Efforts to combat the disease through breeding resistant hybrids and vigilant management underscore ongoing challenges in preserving this genus amid its vulnerability to vascular wilt pathogens.

Description and Morphology

Physical Characteristics

Elms comprise or trees in the genus Ulmus, typically reaching heights of 15 to 40 meters, with trunk diameters up to 2 meters in mature specimens, though sizes vary by and . Many exhibit a vase-shaped or umbrella-like crown formed by ascending branches that spread outward, supported by a straight central trunk. Twigs are slender, often pubescent when young, and some develop corky wings on younger branches. The bark is generally gray to dark brown, developing deep furrows with intersecting ridges or diamond-shaped patterns as the tree ages, providing a distinctive textured appearance. Inner bark in certain , such as slippery elm (U. rubra), is mucilaginous when moistened. Leaves are alternate, simple, ovate to elliptic, measuring 5 to 15 cm in length, with doubly serrate margins and characteristically asymmetrical bases, a diagnostic trait for the ; the upper surface is rough to the touch in many , while the underside may be pubescent. Flowers are small and inconspicuous, typically greenish-red or purplish, borne in drooping clusters or fascicles of 3 to 10, emerging in late winter or early spring before leaf expansion; they are wind-pollinated and lack petals, consisting of a calyx and 4-9 stamens. The is a single-seeded samara, flat and elliptic to obovate, 1 to 2 cm long, with a papery wing surrounding the , maturing in spring and dispersing by .

Growth and Lifecycle

Elms (genus Ulmus) exhibit a lifecycle characterized by rapid juvenile growth, early reproductive maturity, and potential longevity exceeding two centuries in undisturbed conditions. Flowering occurs in early spring, typically 2-3 weeks before leaf flush, with inconspicuous wind-pollinated flowers producing samaras that mature and disperse within weeks. For U. americana, seed production commences as early as age 15, becoming abundant after age 40, with trees remaining productive up to 300 years. Across , reproductive maturity varies from 8 years in U. pumila to 30-40 years in U. glabra. Samaras, containing single seeds, are wind-dispersed up to 0.4 km and exhibit minimal ; is and rapid, peaking in 6-12 days for U. americana under alternating temperatures of 20°C night/30°C day, with viability persisting on flooded soils for a month. Most Ulmus species require no pretreatment, though U. americana and U. rubra benefit from 2-3 months cold stratification; full may extend to 60 days. Seedlings establish best in partial (one-third full exposure) initially, transitioning to full sun after 1-2 years, and develop slowly in saturated or shaded soils. Juvenile elms demonstrate vigorous growth, with U. americana achieving 30-38 m height and 122-152 cm on optimal sites, classified as fast to moderate overall. Rock elm (U. thomasii) seedlings reach 27 cm in 5 years and 52 cm in 10 years post-planting. Vegetative propagation via stump sprouting is common in young trees, with root suckering in dense stands, enabling persistence post-disturbance. Maturity brings canopy dominance in early-successional habitats, though growth slows in sapling-to-pole stages for species like U. rubra. Lifespan ranges from 175-200 years typically, with exceptional individuals surpassing 300 years; factors like site quality and absence dictate duration, as elms invest in height and breadth for light capture before prioritizing . Annual cycles involve winter , spring flush and , summer vegetative expansion, and autumn , with seed crops recurring every 2-4 years. accelerates under stress, but healthy specimens sustain multi-century lifecycles through iterative and seeding.

Taxonomy and Phylogeny

Classification and Species

The genus Ulmus L., commonly known as elm, is placed in the family Mirbel, order , class Magnoliopsida (flowering plants), phylum Tracheophyta, kingdom Plantae. This classification reflects molecular and morphological analyses confirming Ulmaceae's position within , distinct from related families like or based on floral and fruit characteristics such as apetalous flowers and samara fruits. The encompasses 20 to 45 of mostly trees, with the range arising from ongoing taxonomic revisions driven by extensive hybridization, which blurs boundaries through intermediate forms and . The database accepts 37 as of its latest compilation, prioritizing nomenclatural stability and phylogenetic evidence from . are often divided into subgenera such as Ulmus (with typically asymmetrical bases) and Orya (symmetrical leaves), though sectional delimitations remain debated due to in traits like and bark texture. Key accepted species include:
  • Ulmus americana L. (American elm), characterized by large, vase-shaped crowns and serrated leaves up to 15 cm long.
  • Ulmus rubra Muhlenberg (slippery elm), distinguished by mucilaginous inner bark and asymmetrical leaves.
  • Ulmus thomasii Sarg. (rock elm), with corky wings on branches and doubly serrated leaves.
  • Ulmus alata Michaux (winged elm), featuring prominent corky ridges on twigs.
  • Ulmus crassifolia Nuttall (cedar elm), noted for small, thick leaves and early autumn coloration.
  • Ulmus glabra Hudson (wych elm), with large, rounded leaves and no corky wings.
  • Ulmus minor J. Miller (field elm), exhibiting variable leaf shapes and tolerance to wet soils.
  • Ulmus procera Salisbury (English elm), often clonal via root suckering with upright branches.
  • Ulmus pumila L. (Siberian elm), fast-growing with small leaves and invasive potential outside native range.
  • Ulmus parvifolia Jacquin (Chinese elm), semi-evergreen with exfoliating bark and heat tolerance.
Hybrids such as Ulmus × hollandica Mill. (Dutch elm, U. minor × U. glabra) are common in cultivation and natural settings, complicating field identification without genetic analysis. Taxonomic treatments emphasize morphology (e.g., samara notch position) and venation patterns for differentiation, as these traits show less plasticity than growth form.

Evolutionary History

The Ulmus first appears in the record during the early epoch, approximately 50 million years ago, represented by leaf and impressions from deposits in . These early fossils indicate an Asian origin for the genus, with subsequent dispersal to evidenced by middle to late Eocene specimens of leaves and fruits from northwestern regions. The broader Ulmaceae family, encompassing Ulmus, exhibits a more ancient lineage, with macrofossils from the early (~66-56 million years ago) across the and molecular clock estimates placing its diversification in the (ca. 110-125 million years ago). Diversification of Ulmus species accelerated during the epoch (23-5 million years ago), a period marked by the proliferation of temperate forests amid global cooling and tectonic uplift in and . Fossil fruits and woods from Miocene strata in Southwest and other sites document this radiation, with diversity peaking before a Pliocene-Quaternary decline linked to , glaciation, and . Biogeographic centers of emerged in eastern Asia (particularly ) and the , reflecting vicariance and long-distance dispersal facilitated by winged samaras. Molecular phylogenetic analyses, incorporating phylogenomics, chloroplast genomes, and nuclear markers, delineate Ulmus into clades aligned with continental distributions—Eurasian and North American— with intraspecific divergences often dated to the late Miocene-Pliocene transition (ca. 5-6 million years ago). These studies corroborate of adaptive radiations in response to climatic oscillations, including the of traits like asymmetric leaves and corky bark for temperate resilience, though ongoing refinements in dating and sampling continue to refine interclade relationships.

Distribution and Ecology

Geographic Range

The genus Ulmus encompasses 20–40 species native predominantly to the temperate zones of the , with concentrations in and extensions into ; some taxa extend into subtropical and montane tropical areas. Highest species diversity occurs in eastern , particularly , where endemics such as U. chenmoui and U. prunasepala are restricted to specific provinces, alongside widespread species like U. parvifolia (native to central and southern , Korea, , , and ). In western and central , U. pumila spans a vast area from eastern and westward to the , including northern , , and northern , thriving in arid steppes and river valleys. Himalayan species such as U. wallichiana occupy elevations from 1,000 to 3,300 meters across , Pakistan, , and . In , three primary native dominate: Ulmus glabra (wych elm), with the broadest distribution from eastward to the and from the south to the ; U. minor (field elm), common in western and including the Mediterranean basin; and U. laevis (European white elm), centered in along rivers and floodplains from to . These favor riparian and woodland habitats, though their ranges have been fragmented by historical and disease. North American natives are concentrated in the eastern and and adjacent , with U. americana (American elm) extending from and westward to and , southward to and , often in floodplains and bottomlands. Other eastern species include U. rubra (slippery elm) in similar ranges from to northern and , and U. thomasii (rock elm) in northern hardwoods from to and south to . Southern extensions feature U. crassifolia (cedar elm) in and , while Mexican species like U. mexicana occur in northeastern and adjacent southwestern U.S. borders. Isolated populations and hybrids reflect post-glacial migrations, but many ranges overlap in riparian zones.

Habitat and Environmental Adaptations

Elms of the Ulmus primarily inhabit temperate regions across the , favoring riparian zones, floodplains, river valleys, and moist forest edges where fertile soils support rapid growth. Species such as U. americana (American elm) commonly occupy bottomlands and terraces with clay or silty-clay loams, achieving medium growth on wetter sites and optimal development on well-drained uplands. In , U. minor (field elm) and U. glabra (wych elm) associate with lowland woodlands and alluvial soils, often along watercourses that provide seasonal moisture. These habitats reflect elms' ecological role in stabilizing sediments and filtering runoff in dynamic fluvial environments. Elms demonstrate versatile adaptations to soil variability, tolerating textures from clay and to , and pH extremes including alkaline conditions, as seen in U. parvifolia (Chinese elm). Shallow systems in wet soils enable widespread lateral spread for anchorage, conferring windfirmness despite reduced depth, a trait prominent in U. americana. Many species exhibit moderate flood tolerance, enduring infrequent, short-duration inundation—up to several weeks in U. americana—via physiological mechanisms that mitigate anaerobic stress in saturated roots. However, prolonged waterlogging can impair growth, underscoring limits to this . Drought resistance varies across species and populations, with U. americana and U. parvifolia showing reasonable tolerance through efficient water-use strategies and deep rooting in drier contexts, though U. minor proves vulnerable, experiencing heightened stress and susceptibility to secondary pathogens under deficit conditions. Climatic adaptations include local for hardiness; genotypes from northern latitudes in U. americana exhibit greater mid-winter tolerance, reflecting evolutionary tuning to regional gradients via traits like enhanced freezing resistance in tissues. Such plasticity supports elms' persistence in transitional zones but highlights species-specific constraints amid intensifying environmental variability.

Reproduction and Population Dynamics

Elms in the genus Ulmus primarily reproduce sexually through -pollinated flowers that emerge in early spring before leaf expansion, with most being monoecious and producing both staminate and pistillate flowers in small clusters. occurs exclusively via anemophily, with effective dispersal limited to short distances averaging around 50 meters, as demonstrated in studies of U. minor. Following , female flowers develop into single-seeded samaras—winged achenes—that ripen within a few weeks and are primarily dispersed by over short ranges, typically less than 30 meters in like U. laevis. Samaras exhibit orthodox storage behavior in some species, maintaining viability for up to 5 years at low temperatures (1-3°C), though rates decline thereafter, with fresh seeds showing 90-100% viability in tests of U. thomasii. requires cold stratification in many cases and occurs rapidly upon dispersal in spring, but seeds remain viable only for days to weeks post-maturity unless conditions are optimal, limiting long-term persistence. establishment can be abundant under favorable moist, shaded conditions, yet success is constrained by herbivory, competition, and pathogen exposure. Vegetative reproduction via root suckering is prevalent, particularly in European species like field elm (U. minor), where adventitious shoots arise from lateral roots, enabling clonal spread and persistence of genotypes even after bole death. This mode forms dense thickets and maintains population structure through ramets connected to a shared , with suckers capable of developing into mature trees. Population dynamics of elms are characterized by episodic regeneration cycles influenced heavily by Dutch elm disease (Ophiostoma novo-ulmi), which has reduced mature tree densities by over 90% in affected North American and European landscapes since the 1970s, shifting reliance toward juvenile cohorts from suckers and seedlings. In unmanaged stands, such as those of wych elm (U. glabra), disease-induced mortality promotes clonal proliferation via suckering, leading to reduced genetic diversity over decades as sexual recruitment declines relative to vegetative regrowth. This results in patchy distributions with high sucker densities near parent clones but vulnerability to synchronous die-off, as ramets inherit susceptibility; however, heterogeneous landscapes foster pockets of persistence through variable disease pressure and occasional seedling input from surviving reproductives. Restoration efforts emphasize propagating disease-resistant genotypes to bolster sexual reproduction and diversify populations beyond clonal dominance.

Interactions with Other Organisms

Elms are wind-pollinated (anemophilous), producing small, clustered flowers in early spring that release large quantities of lightweight dispersed by air currents, enabling cross- across populations. Although animal vectors are not essential, bees and select native bees forage on this , utilizing it as an early-season protein source amid limited floral alternatives. Species in the genus Ulmus form mutualistic associations with arbuscular mycorrhizal fungi (Glomeromycotina), which colonize to facilitate and other acquisition from , enhancing seedling establishment and drought resilience. Inoculation experiments with have shown AMF symbiosis increases biomass and alleviates salt stress effects, underscoring its role in soil microbial networks. Urban studies of hybrid elms like U. × hollandica confirm persistent AMF communities along urbanization gradients, linking fungal diversity to . Foliage and twigs support diverse non-pest communities, with over 500 North American species utilizing elms for feeding, , or ; notable examples include elm-specialist lepidopteran larvae such as the double-toothed prominent moth (Phlogophora iris). Leaves, characterized by low carbon-to-nitrogen ratios and elevated , are palatable to browsing vertebrates, historically harvested as in and . Samaras, maturing abundantly in , function as soft mast, furnishing essential nutrition for granivorous birds (e.g., cardinals) and small mammals (e.g., squirrels, chipmunks) during seasonal food scarcity. In habitats, this seed crop bolsters early breeding populations of seed-dependent . The branching offers nesting substrates for songbirds and refuge for canopy-dwelling arthropods, while submerged logs in riparian zones provide durable woody for aquatic macroinvertebrates due to elm's decay resistance. In temperate forests, early leaf-out supports migratory passerines during spring stopovers, integrating elms into broader trophic dynamics.

Threats and Pathogens

Dutch Elm Disease

Dutch elm disease (DED) is a lethal vascular primarily affecting elm trees in the Ulmus, caused by the ascomycete fungi Ophiostoma ulmi and the more virulent Ophiostoma novo-ulmi. The fungi invade the tree's vessels, producing mycelia and toxins that block water conduction, leading to and eventual death. American elm (U. americana) is highly susceptible, while species like Siberian elm (U. pumila) show greater tolerance. Symptoms typically emerge in , beginning with and yellowing of leaves on one or more outer crown branches, progressing to browning and curling while leaves remain attached to stems—a known as "flagging." Internal diagnostic signs include dark streaking in the sapwood under the bark, visible upon peeling, confirming fungal invasion. Infected trees may die within a single growing season if symptoms appear early, or decline over 1–3 years if infection occurs later. The pathogens spread via elm bark beetles (Scolytus spp.), which carry fungal spores from overwintering galleries in infected wood to feeding sites on healthy twigs, inoculating the tree during the adult beetle's spring emergence. Root grafts between adjacent elms enable belowground transmission over distances up to 30 meters, amplifying local outbreaks. Human activities, such as moving untreated or logs, further disseminate the . Originating in , DED was first documented in northwest around 1910, with significant research in the from 1914–1919 identifying the fungal cause. It reached the in the 1930s via imported elm logs from , sparking epidemics that spread eastward from and westward to the by 1973. A second, more destructive wave in the 1960s–1970s, driven by O. novo-ulmi, intensified mortality across and . The disease has killed over 40 million American elms in the U.S. alone, representing more than 75% of urban and populations in affected areas, and approximately 30 million elms in the UK during the outbreak. European losses from the initial reached 10–40% in multiple countries by the . Ecologically, it altered forest canopies, reduced dependent on elms, and reshaped urban landscapes where elms were dominant street trees. Management relies on sanitation—prompt removal and destruction (burning, chipping, or debarking) of infected trees to eliminate breeding sites and break transmission cycles—which is the most effective and cost-efficient strategy when implemented community-wide. Trenching to depths of 1–1.5 meters severs root grafts between healthy and infected trees. Insecticides targeting s, such as sprays or systemic injections, provide short-term suppression but require annual reapplication. Preventive injections (e.g., or thiabendazole) can protect high-value trees for 1–3 years but are expensive, labor-intensive, and ineffective for trees with >5% canopy infection. Planting resistant elm cultivars or hybrids offers long-term resilience, though no method eradicates the entirely.

Other Diseases and Phytoplasma

Verticillium wilt, caused by the soilborne fungi Verticillium dahliae and V. albo-atrum, affects elms by invading the vascular system, leading to and yellowing of leaves on individual branches or the entire , often with vascular discoloration appearing as brown streaks in the sapwood. Symptoms typically emerge in spring or early summer, progressing to branch dieback, and infected trees may survive but remain weakened, with mortality rates varying by species and environmental stress. The persists in for years via microsclerotia, and no effective chemical controls exist, though resistant cultivars like certain Asian elms show reduced susceptibility. Canker diseases, induced by fungi such as Neofabraea ulmi or Physalospora ulmi, produce sunken, discolored lesions on branches and trunks, often following wounds, resulting in dieback and gum exudation. These infections are more prevalent in stressed trees and can girdle branches, but they are generally less lethal than vascular wilts, managed through and . Elm yellows, also known as elm phloem necrosis, represents a severe disease caused by 'Candidatus Phytoplasma ulmi', a wall-less, -limited bacterium that disrupts transport by the inner , which turns yellowish-brown to caramel-colored. Symptoms initiate in mid- to late summer with , epinasty (downward curling) of leaves, premature defoliation, and branch dieback, culminating in tree death within one to two years for susceptible species like . The pathogen is vectored primarily by the elm leafhopper (Scaphoideus luteus in , Scaphytopius luteolus in ), with transmission occurring during feeding on sap, and root grafts between trees facilitating spread. First documented in the United States in , elm yellows has caused significant mortality in the eastern U.S. and parts of , though underreported due to symptom overlap with . No curative treatments are available; confirmed infections require immediate tree removal and destruction to prevent vector transmission, with PCR-based diagnostics essential for accurate identification given the pathogen's quarantine status in the . Asian elm species exhibit partial resistance, informing breeding efforts.

Insect and Vertebrate Pests

Elm trees face damage from various pests, primarily defoliators and sap feeders, which can weaken trees through foliage loss or physiological stress. The (Xanthogaleruca luteola), an from , is a major defoliator; its larvae skeletonize leaves by feeding on the lower surface between veins, causing brown, lacy foliage that may drop prematurely, while adults chew irregular round holes. Repeated defoliation over years reduces tree vigor, branch dieback, and growth, though single-year outbreaks rarely kill mature trees. Sap-feeding , including and scales, induce curling, , or from honeydew excretion. such as the woolly elm aphid (Eriosoma spp.) and elm sack gall aphid (Tetraneura ulmi) cause leaf rolling or pouch-like , with limited direct damage but potential for secondary issues like reduced . European elm scale (Eulccanium tiliae) and calico scale suck sap from twigs and branches, leading to yellowed leaves, premature drop, and dieback in heavy infestations. Bark beetles (Scolytus spp.), including the European and native elm bark beetles, bore into , creating galleries that girdle branches and weaken trees, though their impact is often compounded by disease transmission elsewhere documented. Other occasional pests include Japanese beetles, , and leafminers, which contribute to foliage loss but are not elm-specific. Vertebrate pests of elms are less commonly reported and typically affect young or stressed trees through or , with deer ( spp.) occasionally consuming foliage or rubbing antlers against bark, causing wounds that invite secondary infections. Squirrels may chew bark or consume seeds, but such damage remains minor compared to insect impacts in most ecosystems. Overall, vertebrate herbivory does not pose a widespread threat to established elms.

Abiotic Stressors Including Climate Impacts

Elms exhibit varying degrees of tolerance to abiotic stressors, with being a primary limiter of growth and survival across . Riparian field elm () demonstrates sensitivity to , particularly under spring dry-warm conditions and reduced river flows, leading to decreased radial growth and elevated wood δ¹³C values indicative of stress. Saplings of this respond acutely to short-term , with declines in , net , and stomatal conductance as key physiological indicators. Siberian elm () varieties from arid regions show intraspecific variation in , assessed through morphological, physiological, and transcriptional responses, where provenances from severe zones maintain higher survival rates via enhanced osmotic adjustment and antioxidant activity. Such stress often exacerbates biotic vulnerabilities, as -deficient or compacted soils increase susceptibility to in multiple Ulmus . Flooding represents another significant stressor, though some elms adapted to margins exhibit partial resilience. U. minor displays functional adjustments to prolonged flooding, including altered aeration and uptake, but prolonged submersion reduces overall vigor and predisposes trees to secondary decay. American elm (Ulmus americana) in forests can endure periodic inundation from storms and ice-melt, contributing to their role in stabilizing riparian ecosystems, yet excessive or prolonged flooding disrupts and health. Soil-related abiotic factors, including compaction, , and nutritional imbalances, further compound these effects; for instance, heavy metal toxicity tolerance varies by , with no universal resistance across metals. Mechanical injuries from , , or also impair survival, as documented in European elm populations where such events account for notable mortality alongside . Temperature extremes influence elm hardiness, with evidence of local climatic adaptation. Genotypes of U. americana from northern latitudes exhibit superior cold tolerance, as measured by electrolyte leakage assays, reflecting evolutionary adjustments to regional winters rather than broad . Heat stress, often coupled with , impairs , though select cultivars in national trials demonstrate moderate resilience to combined thermal and water deficits. Climate change amplifies these stressors through intensified droughts, erratic flooding, and shifting temperature regimes, potentially contracting elm ranges in vulnerable areas. Projections indicate that drought-sensitive species like U. minor face heightened decline risks in Mediterranean and riparian zones, where reduced precipitation and warmer soils synergize with biotic threats to lower population viability. In North America, U. americana may benefit from habitat specificity in cooler, mesic environments but faces challenges from expanded extreme weather, including frost events outside typical hardening periods. Breeding programs prioritize abiotic tolerance screening, with trials evaluating Ulmus hybrids for performance under simulated climate scenarios, emphasizing genotypes that sustain growth amid projected increases in aridity and thermal variability. Overall, while elms possess adaptive traits like deep rooting for water access, unmitigated climate shifts could override these, underscoring the need for provenance-based restoration to match local abiotic profiles.

Cultivation and Breeding

Traditional Cultivation Practices

In , species such as field elm (Ulmus minor) and English elm (Ulmus procera) were traditionally cultivated for hedgerows through via root suckers, enabling rapid establishment of dense barriers for and field boundaries from medieval periods onward. This suckering habit facilitated natural spread without extensive planting, with trees integrated into mixed hedgerows alongside shrubs like hawthorn. Hedgerows were laid or pleached periodically to maintain structure, providing both timber and leaf fodder through practices. Seed propagation was the primary method across elm species, particularly for and ornamental planting. Samaras, ripening in spring or fall, were collected by sweeping from the ground or stripping from branches shortly after dispersal to avoid viability loss. For species like American elm (U. americana), seeds required cold stratification at 5°C for 2-3 months to break , followed by shallow (0-6.4 mm depth) in nurseries, yielding densities of about 5 seedlings per square meter. One-year-old nursery stock was then field-planted for shade, windbreaks, or avenues, a common practice in during the 18th and 19th centuries. Management involved periodic or to harvest wood and foliage sustainably, with cuts promoting resprouting for in summer "" growth. These techniques, rooted in pre-20th-century European , supported elm's role in systems yielding tough, elastic timber for tools, wheels, and furniture.

Cultivars and Hybrids

Numerous cultivars and hybrids of Ulmus species have been developed through programs since the 1930s to counter the impacts of (DED), prioritizing resistance derived from Asian species, vase-shaped growth forms suitable for urban planting, and tolerance to environmental stresses. In , programs at institutions like and the USDA have focused on interspecific hybrids, crossing susceptible native species such as U. americana with resistant Asian elms like U. davidiana var. japonica and U. parvifolia, yielding clones with superior vascular defenses against the Ophiostoma novo-ulmi . European efforts, including Italy's program initiated in 1975 by the Institute of , have similarly produced hybrids from local U. minor and U. glabra crossed with Asian , though field trials indicate variable long-term survival rates influenced by local strains and climate. Asian hybrids dominate resistant selections due to evolutionary co-adaptation with DED-like pathogens, with North American cultivars like 'Accolade' (U. davidiana var. japonica 'Morton', selected 1990) demonstrating over 90% foliage retention post-inoculation in trials and a mature height of 40-50 feet with upright branching. 'Sapporo Autumn Gold' (U. pumila × U. davidiana var. japonica, released 1970s by the Sapporo Research Station) offers rapid growth to 40 feet, golden fall color, and consistent DED resistance in USDA zones 3-7, though it may suffer from Siberian elm's susceptibility to elm leaf beetle. Other notable hybrids include 'Frontier' (U. hybrids, USDA breeding, 1970s), which reaches 40-50 feet with a broad canopy and high resistance confirmed in multi-year field tests, and 'Commendation' (Ulmus 'Morton Stalwart', Morton Arboretum, 2000s), valued for its stalwart trunk and resistance to both DED and elm yellows. Pure U. americana cultivars exhibit tolerance rather than full resistance, with selections like 'Valley Forge' (released 1995 by USDA) showing less than 10% canopy loss in artificial inoculations and a classic vase form maturing at 50-70 feet, derived from progeny of naturally surviving trees in Ohio. 'New Harmony' (U. americana, USDA, 1995) similarly tolerates DED with minimal wilting in zone 3-9 trials but requires vigilant pruning to prevent vector spread. European hybrids such as 'Columella' (U. minor × U. glabra, Dutch breeding, 1980s) form narrow, columnar trees to 30 meters with moderate DED resistance, while 'Lutece' (Ulmus 'Nanguen', Dutch, 2000s) provides upright growth and enhanced tolerance in complex hybrid lineage. Despite these advances, no cultivar achieves complete immunity, and efficacy depends on early detection and integrated management, as evidenced by ongoing monitoring in arboreta where some hybrids show 20-30% infection rates under high disease pressure.
CultivarParentageKey TraitsOrigin
AccoladeU. davidiana var. japonicaVase-shaped, 40-50 ft, >90% DED resistance,
Valley ForgeU. americanaVase form, 50-70 ft, DED toleranceUSDA,
FrontierU. hybrids (incl. U. parvifolia)Broad canopy, 40-50 ft, high resistanceUSDA,
LuteceComplex Ulmus hybridUpright, moderate resistance

Recent Resistance Breeding Efforts

Efforts to breed elm resistance to (DED) have intensified since the 2010s, focusing on hybridizing susceptible species like with resistant Asian species such as U. pumila and U. parvifolia, alongside screening natural survivors for heritable tolerance. These programs emphasize empirical inoculation trials to quantify resistance, measuring metrics like foliage wilting percentages rather than relying on anecdotal survival. By 2022, controlled crosses from moderately resistant parents yielded three genotypes exhibiting less than 30% average wilting after two years of Ophiostoma novo-ulmi inoculation, outperforming parental lines and indicating polygenic resistance traits amenable to selection. In the United States, the U.S. Forest Service and partners like the Minnesota Invasive Terrestrial Plants and Pests Center have advanced field-based screening since 2020, identifying DED-tolerant U. americana selections from wild populations and propagating them via cloning for multi-site trials. As of August 2025, over a dozen tolerant selections were planted across eight sites in four states to monitor genotype-by-environment interactions, revealing that resistance efficacy varies by regional pathogen strains and climate, with some genotypes showing reduced defense activation in warmer conditions. The Morton Arboretum's program, continuing George Ware's hybrid work, integrates pest resistance screening, producing cultivars like the 2010s-era 'Triumph' (U. 'Morton Glossy') with demonstrated DED tolerance alongside moderate elm leaf beetle resistance. European initiatives, such as Italy's long-term program, have released cultivars like 'Morfeo' (U. 'Morfeo') in the 2010s, validated through repeated inoculations showing superior vascular compartmentalization against fungal invasion compared to susceptible controls. Challenges persist, including unintended trade-offs like heightened susceptibility to secondary stressors in hybrids and the need for diverse to counter evolving , as evidenced by genotype-specific failures in deployment trials. Despite these, of resistant selections has scaled, with over 18 DED-tolerant cultivars available by 2023, four pure U. americana and the rest hybrids, enabling urban and restoration plantings.

Propagation and Management Techniques

Elms are primarily propagated vegetatively to preserve desirable traits such as Dutch elm disease resistance in cultivars, using methods like stem cuttings, root cuttings, and grafting. Softwood stem tip cuttings of Ulmus americana taken in June, treated with indolebutyric acid, root effectively under mist propagation systems. Hardwood cuttings from dormant branches, stored at 35°F for 2-12 weeks, achieve rooting rates when planted in perlite or similar media, often requiring bottom heat and shade during summer. Root cuttings, 2-6 inches long from large-diameter roots, are collected in late fall or winter and induced to sprout indoors before outdoor transplanting, succeeding in species like Ulmus alata. Bench grafting onto seedling rootstocks facilitates clonal multiplication of resistant hybrids, as demonstrated in breeding programs testing for pathogen tolerance. Seed propagation occurs naturally via wind-dispersed samaras maturing in spring, but requires cold stratification at 35-41°F for 30-90 days to break and achieve rates exceeding 50% in controlled settings. However, due to variable susceptibility to diseases, seeds are less favored for commercial cultivation of specific genotypes, with emerging for elite selections like Chinese elm (U. parvifolia), yielding plantlets in 6 months via shoot proliferation on media with cytokinins. Air layering and mound layering supplement field for mature trees, promoting adventitious roots on girdled branches buried in moist . Management emphasizes site selection in well-drained, loamy soils with 5.8-8.0 and full sun exposure to support vigorous growth up to 100 feet in height. Young elms require deep, infrequent watering to establish roots, transitioning to after 2-3 years, while mulching suppresses weeds and conserves moisture without exceeding 3 inches depth to prevent rot. occurs in late fall after leaf drop or early spring before break to minimize flow and entry, focusing on removing co-dominant stems and water sprouts to enhance structural integrity. Fertilization applies balanced NPK formulas sparingly in spring for nutrient-poor sites, avoiding excess that promotes weak growth susceptible to pests. Integrated pest management prioritizes sanitation by promptly removing infected branches and injecting thiabendazole into vascular tissue for suppression, achieving up to 90% efficacy in early-stage infections when applied annually. Trees should be spaced at least 50 feet apart to reduce root grafting transmission of , with monitoring for elm vectors via pheromone traps in urban settings. In forestry contexts, selective thinning maintains canopy diversity, while avoiding mechanical injury to bark preserves natural defenses against fungi. Resistant cultivars like '' demand vigilant scouting, as no technique guarantees immunity amid evolving strains.

Conservation and Restoration

Genetic Conservation Strategies

Genetic conservation strategies for elm species (Ulmus spp.) prioritize ex situ methods to preserve genetic diversity eroded by Dutch elm disease (DED) pandemics, which have caused widespread mortality since the 20th century. These approaches focus on capturing genotypes from remnant populations, including putative resistant individuals, to support future resistance breeding and restoration efforts. In situ dynamic conservation complements ex situ by managing natural populations to facilitate adaptation through natural selection, though ex situ dominates due to DED threats. Ex situ conservation relies on clone banks established via vegetative propagation, such as and softwood cuttings, to replicate and store specific genotypes without . In , the National Programme for the Conservation of Native Elm Genetic Resources, launched in 1987, maintains 441 clones at the Guémené-Penfao nursery, including 205 U. minor, 107 U. × hollandica, 29 U. glabra, and 100 U. laevis. This collection, supplemented by 181 clones from seven countries at Nogent-sur-Vernisson since 2000–2001 under the EU RESGEN-78 , serves as a core resource for hedgerow restoration and evaluation, with a defined core subset of 195 clones. EUFORGEN networks coordinate similar efforts across , emphasizing seed and cutting collections for species like field elm (U. minor), wych elm (U. glabra), and white elm (U. laevis), while recommending habitat protection to bolster small, fragmented populations. Advanced techniques like and enable long-term, space-efficient storage of genetic material, particularly for northern-adapted elms vulnerable to shifts. In , micropropagation of U. laevis and U. glabra uses Driver and Kuniyuki medium with and for shoot initiation, followed by for rooting, though contamination challenges persist. via slow cooling of dormant buds in yields 64% regeneration for U. laevis, comparable to fresh buds, offering pathogen-free, genetically stable preservation superior to traditional methods for recalcitrant species traits. In , cryopreservation covers 100 native and 400 European clones, enhancing viability for decades-long storage. In situ strategies include designating dynamic conservation units, such as France's Val d’Allier (>500 U. laevis individuals) and Ramier de Bigorre (>700 U. laevis), alongside one U. glabra unit at Saint-Pé-de-Bigorre, where silvicultural practices stimulate regeneration despite DED pressure. These units, integrated into EUFORGEN frameworks since the mid-1990s, prioritize protecting seedlings and resprouts in hedgerows and floodplains to maintain evolutionary potential. Challenges include limited true DED resistance in collections and poor natural regeneration in some species, necessitating ongoing evaluation with molecular markers to ensure representativeness and avoid .

Field Restoration Initiatives

Field restoration initiatives for elm species primarily target the reintroduction of and other native elms into forests and natural landscapes decimated by (DED), caused by the fungus Ophiostoma novo-ulmi. These efforts emphasize clonal propagation of survivor trees—those exhibiting natural tolerance—and the planting of hybrids or cultivars with verified resistance, often sourced from long-term breeding programs. In the United States, the Nature Conservancy's Program has led one of the largest such undertakings, planting over 1,900 disease-tolerant American elm ramets across 76 sites in four states since the early 2000s, focusing on riparian and forested areas to restore ecological roles like canopy cover and wildlife habitat. Similar projects in the involve identifying DED-survivor elms in wild populations, them via , and outplanting progeny into forest understories. For instance, collaborations between the Society and regional partners have established test plantations in states like , where resilient clones are trialed in sites such as State Park and Elm Creek Park Reserve to evaluate long-term survival and growth under field conditions. The U.S. Forest Service supports these through systematic screening of survivors and field trials of tolerant selections, aiming to reintegrate elms into diverse woodland ecosystems while monitoring for genotype-by-environment interactions that affect resistance efficacy. In , initiatives like those by the UK's Future Trees Trust conduct field trials of Asian-European hybrids, such as 'Resista' cultivars ('New Horizon' and others), planting them in edges and hedgerows to assess timber quality and DED tolerance over decades. These programs prioritize native or near-native genotypes to minimize genetic pollution, with trials demonstrating survival rates exceeding 90% for select clones after 10–20 years of exposure. Challenges persist, including variable local adaptation and the need for ongoing applications or , but successes in sites like Germany's Eisele nurseries have informed scalable models. Restoration metrics often track metrics like establishment (e.g., >70% in controlled field plots) and canopy , with genetic surveys ensuring diversity to counter evolving strains. Public-private partnerships, such as the U.S. Service's genetic restoration efforts, further advance field planting by propagating resistant U. americana for and floodplain reforestation, as seen in projects replacing ash-dominated stands with elm seedlings post-emerald ash borer decline. These initiatives underscore a shift from urban-centric plantings to broader ecological restoration, with survivor surveys ongoing to identify new candidates for clonal field deployment.

Biotechnology Applications and Debates

Biotechnological applications in elm (Ulmus spp.) primarily focus on techniques for and conservation, as well as to enhance resistance to (DED) caused by Ophiostoma novo-ulmi. protocols enable the clonal propagation of mature, DED-resistant elm genotypes from dormant buds, achieving high multiplication rates under conditions optimized with cytokinins and auxins. has been induced from zygotic embryos of species like and U. glabra, facilitating mass production of uniform planting material for restoration efforts. methods, including slow cooling of shoot tips, support long-term storage of genetic diversity in endangered European elms such as U. glabra and U. laevis. Genetic transformation efforts target DED resistance by introducing antifungal genes, such as the synthetic peptide ESF39A, into American elm (U. americana), resulting in transgenic lines that exhibit reduced vascular streaking and symptom severity in greenhouse inoculations. Researchers at institutions like SUNY ESF have developed protocols for Agrobacterium-mediated gene insertion, confirming stable integration and expression in regenerated plants without disrupting mycorrhizal associations essential for tree health. Genomic resources, including chromosome-level assemblies of U. parvifolia and de novo transcriptomes of resistant U. minor genotypes, identify candidate genes for stress tolerance and pathogen response, informing marker-assisted breeding and CRISPR-based editing strategies. Debates surrounding these applications center on the ecological risks and regulatory barriers to deploying genetically modified (GM) elms in natural ecosystems. Proponents argue that GM trees could restore decimated populations, as lab-tested transgenics show promise against DED, a pathogen that has killed billions of elms since the 1930s. Critics, including voices from the UN Convention on Biological Diversity, highlight uncertainties in gene flow to wild relatives, potential non-target effects on biodiversity, and the need for rigorous long-term field data, given trees' longevity and mobility via pollen and seeds. In the UK, early 2000s trials of GM English elm (U. procera) faced political opposition despite reduced disease symptoms, delaying commercialization and underscoring tensions between biotechnology-driven revival and precautionary environmental principles. While tissue culture remains uncontroversial for clonal propagation, GM releases require balancing empirical evidence of safety against hypothetical risks, with no widespread field deployment as of 2025.

Economic and Practical Uses

Timber Production and Wood Properties

Elm wood exhibits moderate , with specific typically ranging from 0.40 (green) to 0.54 at 12% moisture content across Ulmus . The heartwood is light brown to dark brown, often with a coarse texture and interlocked that enhances shock resistance but complicates splitting and , leading to potential fuzzy surfaces during planing. Mechanically, it is hard and stiff, with Janka values of 810 lbf for English elm (Ulmus procera), 830 lbf for American elm (U. americana), and up to 1,320 lbf for rock elm (U. thomasii); modulus of elasticity averages 9.2 GPa, and modulus of rupture 65 MPa under compression parallel to . These properties confer excellent bending and steam-bending capabilities, making elm suitable for curved furniture components, barrels, and parts historically. It glues and finishes well, though surface preparation is essential due to irregularities, and its supports uses in , boxes, crates, and tool handles, particularly for denser like rock elm. Elm seasons with minimal degrade but is prone to decay if not properly dried, limiting outdoor applications without treatment. Commercial timber production peaked prior to (DED) outbreaks, with elm ranking as the second most important broadleaf species in Britain by volume before the 1970s epidemic, which killed over 25 million trees and collapsed mature stands. In the United States, DED introduction around 1930 similarly devastated urban and rural populations of American elm, reducing harvestable volumes from millions of board feet annually to negligible commercial scales by the late , as infected trees were removed to curb spread. Current production relies on scattered resistant individuals, hybrids, or non-native species like Siberian elm (U. pumila), yielding specialty or veneer rather than bulk timber, with annual U.S. harvests under 1 million board feet as of recent inventories. Restoration efforts prioritize disease-resistant cultivars, but economic viability remains low due to inconsistent supply and competition from more stable hardwoods.

Agricultural and Industrial Applications

Elms have been employed in agricultural systems primarily for roles, such as windbreaks and shelterbelts that mitigate wind erosion, reduce soil desiccation, and enhance crop yields by creating microclimates. In the region of the , American elm (Ulmus americana) was historically a dominant in multi-row windbreaks, often combined with other hardwoods and to provide long-term barriers against , with studies indicating yield increases of up to 20-30% for sheltered crops like and corn. These plantings, established as early as under federal conservation programs, leveraged elm's rapid initial growth and dense foliage for effective wind reduction up to 10-15 times the height of the trees. Certain elm species contribute to in contexts, particularly on marginal lands prone to erosion. Siberian elm (), for instance, has been utilized in arid and semi-arid regions of for dune fixation and , where its extensive root systems and tolerance to poor soils help bind sandy substrates, reducing sediment loss by facilitating vegetation succession and decreasing wind speeds at ground level. In North American applications, elms like winged elm () support naturalized areas and woodland edges in farming landscapes, aiding in slope stabilization and enhancement without requiring intensive management. Industrially, the inner bark of slippery elm () serves as a key raw material for mucilage-based products, harvested sustainably from wild stands in the for extraction of its polysaccharide-rich gum, which is processed into powders, lozenges, and emulsions used in pharmaceuticals as demulcents for soothing irritation in the throat and . Commercial production, peaking in the mid-20th century with annual harvests exceeding 500 tons, incorporates the bark into nutritional supplements and topical ointments, valued for its emollient properties that form a protective upon hydration; however, overharvesting concerns have prompted regulations limiting stripping to trees over 10 inches in diameter. Additionally, elm bark fibers, particularly from slippery and American elms, have been processed into cords and ropes for agricultural tying and netting, exploiting the tensile strength derived from bast processing techniques documented in Native American practices and early industrial milling.

Fodder, Biomass, and Other Utilizations

Elm leaves and young branches have historically served as for in regions where elms are native or naturalized, particularly in and . Species such as Ulmus glabra (wych elm) and Ulmus wallichiana (Himalayan elm) provide nutritious foliage that ruminants like cattle, sheep, and goats consume, with leaf meal from U. wallichiana demonstrating potential as a protein supplement in broiler diets at up to 10% inclusion without adverse effects on growth performance. In traditional European , pollarded elms supplied leaves for winter , enhancing milk quality and animal dental health due to the foliage's abrasive texture. Inner bark from young elms has also been fed to pigs, horses, and calves, offering a digestible feed during shortages. These uses persist in some silvopastoral systems, though modern nutritional analyses emphasize balancing elm with other feeds to avoid potential anti-nutritional factors like . Elm wood, particularly from fast-growing species like Siberian elm (Ulmus pumila), exhibits properties suitable for biomass energy production, including high calorific value and gasification potential comparable to other hardwoods. Studies on U. pumila indicate its biomass yields energy efficiently via thermochemical conversion, with low ash content facilitating combustion or pyrolysis for heat and electricity. While not a primary commercial biomass crop, invasive elm populations in North America present opportunities for harvesting residues for biofuel, potentially mitigating spread while generating renewable energy; for instance, samaras from Siberian elm have been explored as a supplementary feedstock for sustainable bioenergy. Empirical data from gasification trials confirm elm's viability in mixed woody feedstocks, though scalability depends on local availability and disease resistance. Beyond fodder and biomass, elm bark—especially inner bark from slippery elm (Ulmus rubra)—finds use in for its content, which forms a soothing gel when hydrated, aiding conditions like sore throats and gastrointestinal irritation. Harvested sustainably from wild or cultivated trees, the bark's demulcent properties stem from that coat mucous membranes, with historical applications including topical wound treatment and nutritive porridges. Other non-timber applications include emergency human food from boiled leaves or bark meal, as documented in historical European famines, though such uses are limited by and nutritional completeness. Conservation concerns arise from overharvesting U. rubra for markets, prompting calls for cultivated alternatives to preserve wild stocks.

Cultural and Symbolic Roles

Historical and Notable Specimens

The Washington Elm in , was an American elm () traditionally associated with George Washington's assumption of command of the Continental Army on July 3, 1775; the tree, estimated to have germinated in the 1720s or 1730s, stood for approximately 210 years before falling in 1923. Descendant trees propagated from cuttings have been planted at sites including the campus. Another Washington Elm, located near the U.S. Capitol in , was reportedly planted under George Washington's direction and survived until 1948. In , the Constitution Elm (U. americana) provided shade for delegates drafting the state's first constitution during the summer of 1816, when heat made indoor work untenable; this massive tree succumbed to in 1925, but its trunk was preserved in a monument. The Great Elm on , also an U. americana, predated European settlement and stood for over 200 years as a site for public hangings, civic gatherings, and military musters before being felled on February 15, 1876, due to decay. In Europe, the Beauly Elm, a wych elm (Ulmus glabra) at Beauly Priory in the Scottish Highlands, was documented in medieval records and estimated at over 800 years old, making it one of the continent's oldest known specimens until it collapsed in January 2023 from Dutch elm disease. Sapling replacements derived from its lineage were planted at the site in April 2024. A surviving American elm on the Smithsonian grounds in Washington, D.C., planted around the 1860s—predating the adjacent museum by decades—continues to thrive as of 2022, having endured urban development and disease pressures. ![U. americana, Dufferin St., Toronto, c. 1914][float-right]
Many historic elms, once ubiquitous in urban and ceremonial landscapes, were decimated by Dutch elm disease outbreaks starting in the 1930s, reducing North American populations by over 90% in some regions, though resistant cultivars and conservation efforts have preserved genetic lineages from notable trees.

Representations in Art, Literature, and Mythology

In mythology, the elm has been symbolically tied to the and transitions between life and death. Celtic traditions associate elms with elves guarding burial mounds and aiding passage to the , reflecting their role as liminal guardians. sources similarly link elms to , with plane and elm saplings used in sacred groves near the underworld's entrance, as described in Homeric and later classical texts. This recurring motif underscores the tree's perceived connection to mortality, evidenced by its wood's historical use in coffins due to durability and symbolic resonance. In literature, elms frequently embody themes of support, melancholy, and human emotion. The classical "elm and vine" topos, originating in Catullus' Carmen 62 (c. 84–54 BCE) and echoed in Virgil's Georgics (29 BCE) and Ovid's works, depicts the elm as a sturdy husband propping the clinging vine-wife, symbolizing marital interdependence rooted in agricultural practice. This image persisted into English Renaissance drama, as in Shakespeare's Titus Andronicus (c. 1594), where a character invokes it to convey spousal unity amid strife: "You are an elm, my husband, I a vine / Whose weakness, married to thy stronger state, / Makes me with thy strength to communicate." Modern examples include Sylvia Plath's poem "Elm" (1965), which anthropomorphizes the tree as a prophetic, anguished presence tormented by visions, drawing on its form to explore psychological fragmentation. Representations in visual art often highlight elms' aesthetic grandeur and textural details in natural settings. English Romantic painter rendered the species' rugged bark in "Study of the Trunk of an Elm Tree" (c. 1821), an oil sketch executed in that prioritizes empirical observation for lifelike fidelity. In American landscape art, George Inness' "The Elm Tree" (c. 1880) integrates the motif into tonalist compositions, evoking spiritual harmony through the tree's vaulting silhouette against ethereal skies. These works reflect elms' cultural status as emblems of enduring rural beauty prior to widespread 20th-century decline from .

Political and Local Significance

The Liberty Tree, an American elm () located at the corner of Essex and Orange Streets in , served as a central symbol of colonial resistance against British taxation policies. On August 14, 1765, protesters gathered beneath it to oppose the , hanging effigies of tax officials from its branches and igniting a wave of similar actions across the colonies. The tree hosted rallies, celebrations, and flag raisings until British forces felled it in 1775 during of , after which a was erected on its stump to perpetuate the symbol of defiance. This event underscored elms' role in early American political symbolism, representing liberty and collective action against perceived tyranny. In , the Shackamaxon Elm marked the site of William Penn's 1682 treaty with leaders, embodying Quaker ideals of peaceful coexistence and fair dealing between settlers and indigenous groups, though the tree's exact involvement remains tied to foundational myths of the city's origin. Similarly, the Washington Elm in , entered national lore through tradition claiming General assumed command of the Continental Army beneath it on July 3, 1775; while historical evidence for the event is inconclusive, the tree—felled in 1923—shaped public memory and patriotic narratives for over a century. These instances highlight elms' integration into pivotal political moments, often amplified by oral histories despite evidentiary debates. Locally, American elms defined urban landscapes in mid-20th-century , lining streets in cities like and Syracuse where their uniform canopies contributed to aesthetic and economic value, fostering community identity until (DED) epidemics from the 1930s onward prompted aggressive municipal responses. DED's spread, killing tens of millions of trees, spurred cross-jurisdictional policies including , felling mandates, and federal research funding under the U.S. Plant Quarantine Act, though direct eradication efforts remained largely local due to political and logistical hurdles. In , the ' 1921 Forest Law enabled royal decrees for widespread elm condemnation and sanitation, illustrating early state intervention in with lasting precedents for . Preservation controversies persist, as seen in , where debates over removing a century-old American elm in 2021 pitted public attachment against disease risks, reflecting tensions between heritage and practical .

References

  1. https://commons.wikimedia.org/wiki/File:George_Inness_-_The_Elm_Tree_%28ca.1880%29.jpg
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